14 Race and Human Variation
Michael B. C. Rivera
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Learning Objectives
- Illustrate the troubling history of “race” concepts.
- Explain human variation and evolution as the thematic roots of biological anthropology as a discipline.
- Critique earlier “race” concepts based on a contemporary understanding of the apportionment of human genetic variation.
- Explain how biological variation in humans is distributed clinally and in accordance with both isolation-by-distance and Out-of-Africa models.
- Identify phenotypic traits that reflect selective and neutral evolution.
- Extend this more-nuanced view of human variation to today’s research, the implications for biomedical studies, applications in forensic anthropology, and other social/cultural/political concerns.
Humans exhibit biological variation. Humans also have a universal desire to categorize other humans in order to make sense of the world around them. Since the birth of the discipline of biological anthropology, we have been interested in studying how humans vary biologically and what the sources of this variation are. Before we tackle these big problems, first consider this question: Why should we study human variation?

There are certainly academic reasons for studying human variation. First, it is highly interesting and important to consider the evolution of our species (see Chapters 10-13) and how our biological variation may be similar to (or different from) that of other species (e.g., other primates and apes; see Chapters 6 and 7). Second, anthropologists study modern human variation to understand how different biological traits developed over evolutionary time (see Figure 14.1). Suppose we are able to grasp the evolutionary processes that produce the differences in biology, physiology, body chemistry, behaviour, and culture (human variation). In that case, we can make more accurate inferences about evolution and adaptation among our hominin ancestors, complementing our study of fossil evidence and the archaeological record. Third, as will be discussed in more detail later on, it is important to consider that biological variation among humans has biomedical, forensic, and sociopolitical implications. For these reasons, the study of human variation and evolution has formed the basis of anthropological inquiry for centuries and continues to be a major source of intrigue and inspiration for scientific research conducted today.
An even-more-important role of the biological anthropologist is to improve public understanding of human evolution and variation—outside of academic circles. Terms such as race and ethnicity are used in everyday conversations and in formal settings within and outside academia. The division of humankind into smaller, discrete categories is a regular occurrence in day-to-day life. This can be seen regularly when governments acquire census data with a heading like “geographic origin” or “ethnicity.” Furthermore, such checkboxes and drop-down lists are commonly seen as part of the identifying information required for surveys and job applications.
According to professors of anthropology, ethnic studies, and sociology, race is often understood as rooted in biological differences, ranging from such familiar traits as skin colour or eye shape to variations at the genetic level. However, race can also be studied as an “ideological construct” that goes beyond biological and genetic bases (Fuentes Et al. 2019), at different times relating to our ethnicities, languages, religious beliefs, and cultural practices. Sometimes people associate racial identity with the concept of socioeconomic status or position, or they link ideas about race to what passport someone has, how long they have been in a country, or how well they have “integrated” into a population.
Some of these ideas about ethnicity have huge social and political impacts, and notions of race have been part of the motivation behind various forms of racism and prejudice today, as well as many wars and genocides throughout history. Racism manifests in many ways—from instances of bullying between kids on school playgrounds to underpaid minorities in the workforce, and from verbal abuse hurled at people of colour to violent physical behaviours against those of a certain race. Prejudice can be characterized as negative views toward another group based on some perceived characteristic that makes all members of that group worthy of disdain, disrespect, or exclusion (not solely along racial lines but also according to [dis]ability, gender, sexual orientation, or socioeconomic background, for example). According to Shay-Akil McLean (2014), “Racism is not something particular to the United States and race is not the same everywhere in the world. Racial categories serve particular contextual purposes depending on the society they are used in, but generally follow the base logic of the supremacy of one type of human body over all others (ordering these human bodies in a hierarchical fashion).” Choosing which biological or nonbiological features to use when discussing race is always a social process (Omi and Winant 2014). Race concepts have no validity to them unless people continue to use them in their daily lives—and, in the worst cases, to use them to justify racist behaviours and problematic ideas about racial difference or superiority/inferiority. Recent work in anthropological genetics has revealed the similarities amongst humans on a molecular level and the relatively few differences that exist between populations (Omi and Winant 2014).
The role of the biological anthropologist becomes crucial in the public sphere, because we may be able to debunk myths surrounding human variation and shed light, for the nonanthropologists around us, on how human variation is actually distributed worldwide (see Figure 14.1). Rooted in scientific observations, our work can help nonanthropologists recognize how common ideas about “race” often have no biological or genetic basis. Many anthropologists work hard to educate students on the history of where race concepts come from, why and how they last in public consciousness, and how we become more conscious of racial issues and the need to fight against racism in our societies. Throughout this chapter, I will highlight how humans cannot actually be divided into discrete “races,” because most traits vary on a continuous basis and human biology is, in fact, very homogenous compared to the greater genetic variation we observe in closely related species. Molecular anthropology, or anthropological genetics, continues to add new layers to our understanding of human biological variation and the evolutionary processes that gave rise to the contemporary patterns of human variation. The study of human variation has not always been unbiased, and thinkers and scientists have always worked in their particular sociohistorical context. For this reason, this chapter opens with a brief overview of race concepts throughout history, many of which relied on unethical and unscientific notions about different human groups.
Special Topic: My Experiences as an Asian Academic

My name is Michael, and I am a biological anthropologist and archaeologist (Figure 14.2). What strikes me as most interesting to investigate is human biological variation today and the study of past human evolution. For instance, some of my research on ancient coastal populations has revealed positive effects of coastal living on dietary health and many unique adaptations in bones and teeth when living near rivers and beaches. I love talking to students and nonscientists about bioanthropologists’ work, through teaching, science communication events, and writing book chapters like this one. I grew up in Hong Kong, a city in southern China. My father is from the Philippines and my mother is from Hong Kong, which makes me a mixed Filipino-Chinese academic. Growing up, I noticed that people came in all shapes, sizes, and colours. My life is very different now in that I have gained the expertise to explain those differences, and I feel a great responsibility to guide those new to anthropology toward their own understandings of diversity.
Biological anthropology is not taught extensively in Hong Kong, so I moved to the United Kingdom to earn my bachelor’s, master’s, and doctorate degrees. It was fascinating to me that we could answer important questions about human variation and history using scientific methods. However, I did not have many minority academic role models to look up to while I was at university. My department was made up almost exclusively of white westerner faculty, and it was hard to imagine I could one day get a job at these western institutions. While pursuing my degrees, I also remember several instances of my research contributions being overlooked or dismissed. Sometimes professors and fellow students would make racist comments toward Asian scholars (including me) and other Black, Indigenous and researchers of colour, making us greatly uncomfortable in those spaces. When one of us would work up the courage to tell university leaders our experiences of being stereotyped, dismissed, or insulted, we received little support and were further excluded from research and teaching activities. This is a common experience for Black, Indigenous, and other people of colour who pursue biological anthropology, and we face the difficult choice between leaving the field or bearing with such unsafe spaces.
It became important to me at that time to find other academics of colour with whom to share experiences and form community. I feel inspired by all my colleagues who advocate for greater representation and diversity in universities (whether they are minority academics or not). I admire many of my fellow researchers who are underrepresented and do a great job of representing minority groups through their cutting-edge research and quality teaching at the undergraduate and graduate levels. Although I no longer work in the West, it has remained my great hope that those in the West and the “Global North” will continue to improve university culture, and I support any efforts there to welcome all scholars.
My current work is based in Hong Kong, where I am deeply dedicated to helping develop biological anthropology in East and Southeast Asia and promoting research from our home regions on the international scene. The study of anthropology really highlights how we share a common humanity and history. Being somebody who is “mixed race” and Asian likely played a key role in steering me toward the study of human variation. As this chapter hopefully shows, there is a lot to discuss about race and ethnicity regarding the discipline’s history and current understandings of human diversity. Some scientific and technological advancements today are misused for reasons to do with money, politics, or the continuation of antiquated ideas. It is my belief, alongside many of my friends and fellow anthropologists, that science should be more about empathy toward all members of our species and contributing to the intellectual and technological nourishment of society. After speaking to many members of the public, as well as my own undergraduate students, I have received lovely messages from other individuals of colour expressing thanks and appreciation for my presence and understanding as a minority scientist and mentor figure. Anthropology needs much more diversity as well as to make room for those who have traveled different routes into the discipline. All paths taken into anthropology are valid and valuable. I would encourage everyone to study anthropology—it is a field for understanding and celebrating the intricacies of human diversity.
The History of “Race” Concepts
“Race” in the Classical Era

The earliest classification systems used to understand human variation are evidenced by ancient manuscripts, scrolls, and stone tablets recovered through archaeological, historical, and literary research. The Ancient Egyptians had the Book of Gates, dated to the New Kingdom between 1550 B.C.E. and 1077 B.C.E (Figure 14.3). In one part of this tome dedicated to depictions of the underworld, scribes used pictures and hieroglyphics to illustrate a division of Egyptian people into the four categories known to them at the time: the Aamu (Asiatics), the Nehesu (Nubians), the Reth (Egyptians), and the Themehu (Libyans). Though not rooted in any scientific basis like our current understandings of human variation today, the Ancient Egyptians believed that each of these groups were made of a distinctive category of people, distinguishable by their skin colour, place of origin, and even behavioural traits.
Another well-known early document is the Bible, where it is written that all humankind descends from one of three sons of Noah: Shem (the ancestor to all olive-skinned Asians), Japheth (the ancestor to pale-skinned Europeans), and Ham (the ancestor to darker-skinned Africans). Similar to the Ancient Egyptians, these distinctions were based on behavioural traits and skin colour. More recent work in historiography and linguistics suggest that the branches of “Hamites,” “Japhethites,” and “Shemites” may also relate to the formation of three independent language groups some time between 1000 and 3000 B.C.E. With the continued proliferation of Christianity, this concept of approximately three racial groupings lasted until the Middle Ages and spread as far across Eurasia as crusaders and missionaries ventured at the time.

There is also the “Great Chain of Being,” conceived by ancient Greek philosophers like Plato (427‒348 B.C.E.) and Aristotle (384‒322 B.C.E.). They played a key role in laying the foundations of empirical science, whereby observations of everything from animals to humans were noted with the aim of creating taxonomic categories. Aristotle describes the Great Chain of Being as a ladder along which all objects, plants, animals, humans, and celestial bodies can be mapped in an overall hierarchy (in the order of existential importance, with humans placed near the top, just beneath divine beings; see Figure 14.4). When he writes about humans, Aristotle expressed the belief that certain people are inherently (or genetically) more instinctive rulers, while others are more natural fits for the life of a worker or enslaved person. Based on research by biological anthropologists, we currently recognize that these early systems of classification and hierarchization are unhelpful in studying human biological variation. Both behavioural traits and physical traits are coded for by multiple genes each, and how we exhibit those traits based on our genetics can vary significantly even between individuals of the same population.
“Race” during the Scientific Revolution
The 1400s to 1600s saw the beginnings of the Scientific Revolution in Western societies, with thinkers like Nicholas Copernicus, Galileo Galilei, and Leonardo Da Vinci publishing some of their most important findings. While by no means the first or only scholars globally to use observation and experimentation to understand the world around them, early scientists living at the end of the medieval period in Europe increasingly employed more experimentation, quantification, and rational thought in their work. This is the main difference between the work of the ancient Egyptians, Romans, and Greeks and that of later scientists such as Isaac Newton and Carl Linnaeus in the 1600s and 1700s.

Linnaeus is the author of Systema Naturae (1758), in which he classified all plants and animals under the formalized naming system known as binomial nomenclature (Figure 14.5). This system is typological, in that organisms are placed into groups according to how they are similar or different to others under study. What was most anthropologically notable about Linnaeus’s typological system was that he was one of the first to group humans with apes and monkeys, based on the anatomical similarities between humans and nonhuman primates. However, Linnaeus viewed the world in line with essentialism, a problematic concept that dictates that there are a unique set of characteristics that organisms of a specific kind must have and that would remove organisms from taxonomic categorizations if they lacked any of the required criteria.
Linnaeus subdivided the human species into four varieties, with overtly racist categories based on skin colour and “inherent” behaviours. Some European scientists during this period were not aware of their own biases skewing their interpretations of biological variation, while others deliberately worked to shape public perceptions of human variation in ways that established “otherness” and enforced European domination and the subordination of non-European people. The conclusions and claims at which they arrived, consciously or subconsciously, often fit the times they were living through—the so-called Age of Discovery, when the superiority of European cultures over others was a pervasive idea throughout people’s social and political lives. Although much of Eurasia was linked by spice- and silk-trading routes, the European colonial period between the 1400s and 1700s was marked by many new and unfortunately violent encounters overseas (Figure 14.6). When Europeans arrived by ship on the shores of continents that were already inhabited, it was their first meeting with the Indigenous peoples of the Americas and Australasia, who looked, spoke, and behaved differently from peoples with whom they were familiar. Building on the idea of species and “subspecies,” natural historians of this time invented the term race, from the French rasse meaning “local strain.”

Another scientist of the times, Johann Friedrich Blumenbach (1752‒1840), classified humans into five races based on his observations of cranial form variation as well as skin colour. He thus dubbed the “original” form of the human cranium the “Caucasian” form, with the idea that the ideal climate conditions for early humans would have been in the Caucasus region near the Caspian Sea. The key insight Blumenbach presented was that human variation in any particular trait should be more accurately viewed as falling along a gradation (Figure 14.7). While some of his theories were correct according to what we observe today with more knowledge in genetics, they erroneously believed that human “subspecies” were “degenerated” or “transformed” varieties of an ancestral Caucasian or European race. According to them, the Caucasian cranial dimensions were the least changed over human evolutionary time, while the other skull forms represented geographic variants of this “original.” As will be discussed in greater detail later in this chapter, we have genetic and craniometric evidence for sub-Saharan Africa being the origin of the human species instead (see Chapter 13 on the fossil record that places the origins of modern Homo sapiens in north and east Africa). Based on work that shows how most biological characteristics are coded for by nonassociated genes, it is not reasonable to draw links between individuals’ personalities and their skull shapes.

“Race” and the Dawn of Scientific Racism
Between the 1800s and mid-1900s, and contrary to what you might expect, an increased use of scientific methods to justify racial schemes developed in scholarship. Differing from earlier views, which saw all humans as environmentally deviated from one “original” humankind, classification systems after 1800 became more polygenetic (viewing all people as having separate origins) rather than monogenetic (viewing all people as having a single origin). Instead of moving closer to our modern-day understandings of human variation, there was increased support for the notion that each race was created separately and with different attributes (intelligence, temperament, and appearance).
The 1800s were an important precursor to modern biological anthropology as we know it, given that this was when the scientific measurement of human physical features (anthropometry) truly became popularized. However, empirical studies in the 1800s pushed even further the idea that Europeans were culturally and biologically superior to others. While considered one of the pioneers of American “physical” anthropology, Samuel George Morton (1799‒1851) was a scholar who had a large role in perpetuating 1800s scientific racism. By measuring cranial size and shape, he calculated that “Caucasians,” on average, have greater cranial volumes than other groups, such as the Indigenous peoples of the Americas and peoples Morton referred to collectively as “Negros.” Today, we know that cranial size variation depends on such factors as Allen’s and Bergmann’s rules, which give a more likely explanation: in colder environments, it is advantageous for those living there to have larger and rounder heads because they conserve heat more effectively than more slender heads (Beals Et al. 1984). The leading figures in craniometry during the 1800s instead were linked heavily with powerful individuals and wealthy sociopolitical institutions and financial bodies. Theories in support of hierarchical racial schemes using “scientific” bases certainly helped continue the exploitative and unethical trafficking and enslavement of Africans between the 1500s and 1800s.
Morton went on to write in his publication Crania Americana (1839) a number of views that fit with a concept called biological determinism. The idea behind biological determinism is that an association exists between people’s physical characteristics and their behaviour, intelligence, ability, values, and morals. If the idea is that some groups of people are essentially superior to others in cognitive ability and temperament, then it is easier to justify the unequal treatment of certain groups based on outward appearances. Another such problematic thinker was Paul Broca (1824‒1880), after whom a region of the frontal lobe related to language use is named (Broca’s area). Influenced by Morton, Broca likewise claimed that internal skull capacities could be linked with skin colour and cognitive ability. He went on to justify the European colonization of other global territories by purporting it was noble for a biologically more “civilized” population to improve the “humanity” of more “barbaric” populations. Today, these theories of Morton, Broca, and others like them are known to have no scientific basis. If we could speak with them today, they would likely try to emphasize that their conclusions were based on empirical evidence and not a priori reasoning. However, we now can clearly see that their reasoning was biased and affected by prevailing societal views at the time.
“Race” and the Beginnings of Physical Anthropology
In the early 20th century, we saw a number of new figures coming into the science of human variation and shifting the theoretical focus within. Most notably, these included Aleš Hrdlička and Franz Boas.

Aleš Hrdlička (1869‒1943) was a Czech anthropologist who moved to the United States. In 1903, he established the physical anthropology section of the National Museum of Natural History (Figure 14.8). In 1918, he founded the American Journal of Physical Anthropology, which remains one of the foremost scientific journals disseminating bioanthropological research. As part of his work and the scope of the journal, he differentiated “physical anthropology” from other kinds of anthropology: he wrote that physical anthropology is “the study of racial anatomy, physiology, and pathology” and “the study of man’s variation” (Hrdlička 1918). In some ways, although the scope and technological capabilities of biological anthropologists have changed significantly, Hrdlička established an area of inquiry that has continued and prospered for over a hundred years.
Franz Boas (1858‒1942) was a German American anthropologist who established the four-field anthropology system in the United States and founded the American Anthropological Association in 1902. He argued that the scientific method should be used in the study of human cultures and the comparative method for looking at human biology worldwide. One of Boas’s significant contributions to biological anthropology was the study of skull dimensions with respect to race. After a long-term research project, he demonstrated how cranial form was highly dependent on cultural and environmental factors and that human behaviours were influenced primarily not by genes but by social learning. He wrote in one essay for the journal Science: “While individuals differ, biological differences between races are small. There is no reason to believe that one race is by nature so much more intelligent, endowed with great willpower, or emotionally more stable than another, that the difference would materially influence its culture” (Boas 1931:6). This conclusion directly contrasted with the theories of the past that relied on biological determinism. Biological anthropologists today have found evidence that corroborates Boas’s explanations: societies do not exist on a hierarchy or gradation of “civilizedness” but instead are shaped by the world around them, their demographic histories, and the interactions they have with other groups.

The first half of the 1900s still involved some research that was essentialist and focused on proving racial determinism. Anthropologists like Francis Galton (1822‒1911) and Earnest A. Hooton (1887‒1954) created the field of eugenics as an attempt to formalize the social scientific study of “fitness” and “superiority” among members of 19th-century Europe. As a way of “dealing with” criminals, diseased individuals, and “uncivilized” people, eugenicists recommended prohibiting parts of the population from being married or sterilizing these members of society so they could no longer procreate (Figure 14.9). They instead encouraged “reproduction in individual families with sound physiques, good mental endowments, and demonstrable social and economic capability” (Hooton 1936). In the 1930s, Nazi Germany used this false idea of there being “pure races” to highly destructive effect. The need to be protected against admixture from “unfit” groups was their justification for their blatant racism and purging of citizens that fell under their subjective criteria.
It should be noted that eugenicist ideas were popularly discussed and debated in many non-European contexts, as in the U.S., China, and South Africa, too. The Immigration Restriction Act of 1924 was passed in the United States, with the explicit aim of reducing the country’s “burden” of people considered inferior by restricting immigration of eastern European Jews, Italians, Africans, Arabs, and Asians. In the early 1900s, Chinese scientists and politicians showed great interest in eugenic ideologies, which came to dictate decisions in law-making, family life, and the medical field. Noted American anthropologist Ruth Benedict wrote extensively on Japanese culture and society during and after World War II. Her essentialist portrayals of Japanese people were heavily cited in popular discourse at the time. In 1950s South Africa, interracial marriages and sexual relations were banned by law; antimiscegenation became one of the huge focuses of apartheid resistance activists in later years. We still see the continuation of eugenics-based logic today around the world—in exclusionary immigration laws, cases of incarcerated prison inmates being forcibly sterilized, and the persistence of intelligence testing as a form of measuring people’s “fitness” in a society.
Shortly after World War II and the Nazi Holocaust, the full extent of essentialist, eugenicist thinking became clear. Social constructions of race, and the notion that one can predict psychological or behavioural traits based on external appearance, had become unpopular both within and outside the discipline. It was up to those in the field of physical anthropology at the time to separate physical anthropology from race concepts that supported unscientific and socially damaging agendas. This does not mean that there are no physiological or behavioural differences between different members of the human species. However, going forward, a number of physical anthropologists saw human biological variation as more complicated than simple typologies could describe.
“The New Physical Anthropology”

After 1950, focus steered away from the concept of “race” as a unit of variation and toward understanding why variation exists in populations from an evolutionary perspective. This was outlined by those pioneering the “new physical anthropology,” such as Sherwood Washburn, Theodosius Dobzhansky (Figure 14.10), and Julian Huxley, who borrowed this approach from contemporary population geneticists. Whether using genetic or phenotypic markers as the units of study, “groups” or “clusters” of humans differentiated by these became defined as populations that differ in the frequency of some gene or genes. Anthropologists consider what “makes” a population—a group of individuals potentially capable of or actually interbreeding due to shared geographic proximity, language, ethnicity, culture, and/or values. Put another way, a population is a local interbreeding group with reduced gene flow between themselves and other groups of humans. Members of the same population may be expected to share many genetic traits (and, as a result, many phenotypic traits that may or may not be visible outwardly).

Thinking of humans in terms of populations was part of Julian Huxley’s (1942) “Modern Synthesis”—so named because it helped to reconcile two fundamental principles about evolution that had not been made sense of together before (Figure 14.11). As discussed in Chapter 4, Gregor Mendel (1822‒1884) was able to show that inheritance was mediated by discrete particles (or genes) and not blended in the offspring. However, it was difficult for some 19th-century scientists to accept this model of genetic inheritance at the time because much of biological variation appeared to be continuous and not particulate (take skin colour or height as examples). In the 1930s, it was demonstrated that traits could be polygenic and that multiple alleles could be responsible for any one phenotypic trait, thus producing the continuous variation in traits such as eye colour that we see today. Thus, Huxley’s “Modern Synthesis” outlines not only how human populations are capable of exchanging genes at the microevolutionary level but also how multiple alleles for one trait (polygenic exchanges) can cause gradual macroevolutionary changes.
Human Variation in Biological Anthropology Today
Many Human Traits Are Nonconcordant
In our studies of human (genetic) variation today, we understand most human traits to be nonconcordant (Figure 14.12). “Nonconcordance” is a term used to describe how biological traits vary independent of each other—that is, they do not get inherited in a correlative manner with other genetically controlled traits. For example, if you knew an individual had genes that coded for tall height, you would not be able to predict if they are lighter-skinned or have red hair. This is different from earlier essentialist views of human variation: the idea that skin colour could predict one’s brain function or even “temperament” and tendencies toward criminal behaviour.

Human Variation Is Clinal/Continuous (Not Discrete)
Frank B. Livingstone (1928‒2005) wrote: “There are no races, only clines” (1962: 279). A cline is a gradation in the frequency of an allele/trait between populations living in different geographic regions. Human variation cannot be broken into discrete “races,” because most physical traits vary on a continuous or “clinal” basis. One obvious example of this is how human height does not only come in three values (“short,” “medium,” and “tall”) but instead varies across a spectrum of vertical heights achievable by humans all over the world. On the one hand, we can describe human height as exhibiting continuous variation, forming a continuous pattern, but height does not vary according to where people live across the globe and does not exhibit a clinal pattern. On the other hand, skin colour variation between populations does show patterning that fits quite well on to how near or far they are from each other on a world map. This makes a trait like skin colour clinally distributed worldwide. When large numbers of genetic loci for large numbers of samples were sampled from human populations distributed worldwide during the 1960s and 1970s, the view that certain facets of human diversity were clinally distributed was further supported by genetic data.
To study human traits that are clinally distributed, genetic tests must be performed to uncover the true frequencies of an allele or trait across a certain geographic space. One easily visible example of a clinal distribution seen worldwide is the patterning of human variation in skin colour. Whether in southern Asia, sub-Saharan Africa, or Australia, dark brown skin is found. Paler skin tones are found in higher-latitude populations such as those who have lived in areas like Europe, Siberia, and Alaska for millennia. Skin colour is easily observable as a phenotypic trait exhibiting continuous variation.
A clinal distribution still derives from genetic inheritance; however, clines often correspond to some gradually changing environmental factor. Clinal patterns arise when selective pressures in one geographic area differ from those in another as well as when people procreate and pass on genes together with their most immediate neighbours. There are several mechanisms, selective and neutral, that can lead to the clinal distribution of an allele or a biological trait. Natural selection is the mechanism that produced a global cline of skin colour, whereby darker skin colour protects equatorial populations from high amounts of UV radiation; there is a transition of lessening pigmentation in individuals that reside further and further away from the tropics (Jablonski 2004; Jablonski and Chaplin 2000; see Figure 14.13). The ability and inability to digest lactose (milk sugar) among different world communities varies according to differential practices and histories of milk and dairy-product consumption (Gerbault Et al. 2011; Ingram Et al. 2009). Where malaria seems to be most prevalent as a disease stressor on human populations, a clinal gradient of increasing sickle cell anemia experience toward these regions has been studied extensively by genetic anthropologists (Luzzatto 2012). Sometimes, culturally defined mate selection based on some observable trait can lead to clinal variation between populations as well.

Two neutral microevolutionary processes that may produce a cline in a human allele or trait are gene flow and genetic drift (see Chapter 5). The ways in which neutral processes can produce clinal distributions is seen clearly when looking at clinal maps for different blood groups in the human ABO blood group system (Figure 14.14). For instance, scientists have identified an East-to-West cline in the distribution of the blood type B allele across Eurasia. The frequency of B allele carriers decreases gradually westward when we compare the blood groups of East and Southeast Asian populations with those in Europe. This shows how populations residing nearer to one another are more likely to interbreed and share genetic material (i.e., undergo gene flow). We also see 90%‒100% of native South American individuals, as well as between 70%‒90% of Aboriginal Australian groups, carrying the O allele (Mourant, Kopeć, and Domaniewska-Sobczak 1976). These high frequencies are likely due to random genetic drift and founder effects, in which population sizes were severely reduced by the earliest O allele-carrying individuals migrating into those areas. Over time, the O blood type has remained predominant.



Genetic Variation Is Greater Within Group than Between Groups

One problem with race-based classifications is they relied on an erroneous idea that individuals with particular characteristics would share more similar genes with each other within a particular “race” and share less with individuals of other “races” possessing different traits and genetic makeups. However, since around 50 years ago, scientific studies have shown that the majority of human genetic differences worldwide exist within groups (or “races”) individually rather than between groups. Indeed, most genetic variation we see occurs in Africa, and many variants are shared among individuals on all continents (Figure 14.15).
In 2002, a landmark article by Noah Rosenberg and colleagues explored worldwide human genetic variation using an even-greater genetic data set. They used 377 highly variable markers in the human genome and sampled from 1,056 individuals representative of 52 populations. The markers chosen for study were not ones that code for any expressed genes. Because these regions of the human genome were made of unexpressed genes, we may understand these markers as neutrally derived (as opposed to selectively derived) because they do not code for functional advantages or disadvantages. These neutral genetic markers likely reflect an intricate combination of regional founder effects and population histories. Analyses of these neutral markers allowed scientists to identify that 93%‒95% of global genetic differences, referred to as variance, can be accounted for by within-population differences, while only a small proportion of genetic variance (3%‒5%) can be attributed to differences among major groups (Rosenberg Et al. 2002). This research supports the theory that distinct biological races do not exist, even though misguided concepts of race may still have real social and political consequences.
Biological Data Fit Isolation-By-Distance and Out-of-Africa Models
One further note is that the world’s population may be genetically divided into “groups,” “subsets,” “clumps,” or “clusters” that reflect some degree of genetic similarity. These identifiable clusters reflect genetic or geographic distances—either with gene flow facilitated by proximity between populations or impeded by obstacles like oceans or environmentally challenging habitats (Rosenberg Et al. 2005). Sometimes, inferred clusters using multiple genetic loci are interpreted by nongeneticists literally as “ancestral populations.” However, it would be wrong to assume from these genetic results that highly differentiated and “pure” ancestral groups ever existed. These groupings reflect differences that have arisen over time due to clinal patterning, genetic drift, and/or restricted or unrestricted gene flow (Weiss and Long 2009). The clusters identified by scientists are arbitrary and the parameters used to split up the global population into groups is subjective and dependent on the particular questions or distinctions being brought into focus (Relethford 2009).

Additionally, research on worldwide genetic variation has shown that human variation decreases with increasing distance from sub-Saharan Africa, where there is evidence for this vast region being the geographical origin of anatomically modern humans (Liu Et al. 2006; Prugnolle, Manica, and Balloux 2005; see Figures 14.16 and 14.17). Genetic differentiation decreases in human groups the further you sample data from relative to sub-Saharan Africa because of serial founder effects (Relethford 2004). Over the course of human colonization of the rest of the world outside Africa, populations broke away in expanding waves across continents into western Asia, then Europe and eastern Asia, followed by Oceania and the Americas. As a result, founder events occurred whereby genetic variation was lost, as the colonization of each new geographical region involved a smaller number of individuals moving from the original larger population to establish a new one (Relethford 2004). The most genetic variation is found across populations residing in different parts of sub-Saharan Africa, while other current populations in places like northern Europe and the southern tip of South America exhibit some of the least genetic differentiation relative to all global populations (Campbell and Tishkoff 2008).


Besides fitting nicely into the Out-of-Africa model, worldwide human genetic variation conforms to an isolation-by-distance model, which predicts that genetic similarity between groups will decrease exponentially as the geographic distance between them increases (Kanitz Et al. 2018). This is because of the greater and greater restrictions to gene flow presented by geographic distance, as well as cultural and linguistic differences that occur as a result of certain degrees of isolation. Since genetic data conform to isolation-by-distance and Out-of-Africa models, these findings support the abolishment of “race” groupings. This research demonstrates that human variation is continuous and cannot be differentiated into geographically discrete categories. There are no “inherent” or “innate” differences between human groups; instead, variation derives from some degree of natural selection, as well as neutral processes like population bottle-necking (Figure 14.18), random mutations in the DNA, genetic drift, and gene flow through between-mate interbreeding.
Humans Have Higher Homogeneity Compared to Many Other Species
An important fact to bear in mind is that humans are 99.9% identical to one another. This means that the apportionments of human variation discussed above only concern that tiny 0.1% of difference that exists between all humans globally. Compared to other mammalian species, including the other great apes, human variation is remarkably lower. This may be surprising given that the worldwide human population has already exceeded seven billion, and, at least on the surface level, we appear to be quite phenotypically diverse. Molecular approaches to human and primate genetics tells us that external differences are merely superficial. For a proper appreciation of human variation, we have to look at our closest relatives in the primate order and mammalian class. Compared to chimpanzees, bonobos, gorillas and other primates, humans have remarkably low average genome-wide heterogeneity (Osada 2005).
When we look at chimpanzee genetic variation, it is fascinating that western, central, eastern, and Cameroonian chimpanzee groups have substantially more genetic variation between them than large global samples of human DNA (Bowden Et al. 2012; Figure 14.19). This is surprising given that all of these chimpanzee groups live relatively near one another in Africa, while measurements of human genetic variation have been conducted using samples from entirely different continents. First, geneticists suppose that this could reflect differential experiences of the founder effect between humans and chimpanzees. As it has been argued that all non-African human populations descended from a small number of anatomically modern humans who left Africa, it would be expected that all groups descended from that smaller ancestral group would be similar genetically. Second, our species is really young, given that we have only existed on the planet for around 150,000 to 300,000 years. This gave humans little time for random genetic mutations to occur as genes get passed down through genetic interbreeding and meiosis. Chimpanzees, however, have inhabited different ecological niches, and less interbreeding has occurred between the four chimpanzee groups over the past six to eight million years compared to the amount of gene flow that occurred between worldwide human populations (Bowden Et al. 2012).

Recent advances have now enabled the attainment of genetic samples from the larger family of great apes and the evaluation of genetic variation among bonobos, orangutans, and gorillas alongside that of chimpanzees and humans (Prado-Martinez Et al. 2013). Collecting such data and analyzing primate genetic variation has been important not only to elucidate how different ecological, demographic, and climatic factors have shaped our evolution but also to inform upon conservation efforts and medical research. Genes that may code for genetic susceptibilities to tropical diseases that affect multiple primates can be studied through genome-wide methods. Species differences in the genomes associated with speech, behaviour, and cognition could tell us more about how human individuals may be affected by genetically derived neurological or speech-related disorders and conditions (Prado-Martinez Et al. 2013; Staes Et al. 2017). In 2018, a great ape genomic study also reported genetic differences between chimpanzees and humans related to brain cell divisions (Kronenberg Et al. 2018). From these results, it may be inferred that cognitive or behavioural variation between humans and the great apes might relate to an increased number of cortical neurons being formed during human brain development (Kronenberg Et al. 2018). Comparative studies of human and nonhuman great ape genetic variation highlight the complex interactions of population histories, environmental changes, and natural selection between and within species. When viewed in the context of overall great ape variation, we may reconsider how variable the human species is relatively and how unjustified previous “race” concepts really were.
Phenotypic Traits That Reflect Neutral Evolution
Depending on the trait being observed, different patterns of phenotypic variation may be found within and among groups worldwide. In this subsection, some phenotypic traits that reflect the aforementioned patterns of genetic variation will be discussed.

Looking beyond genetic variation briefly, recent studies have revisited biological anthropology’s earlier themes of externally observable traits, such as skull shape. In the last 20 or so years, anthropologists have evaluated the level to which human cranial shape variation reflects the results from genetic markers, such as those used previously to fit against Out-of-Africa models (Relethford 2004) or those used in the apportionment of human variation between and within groups (Lewontin 1972; Rosenberg Et al. 2002). Using larger sample sizes of cranial data collected from thousands of skulls worldwide and a long list of cranial measurements, studies demonstrate a similar decrease in variation with distance from Africa and show that a majority of cranial variation occurs within populations rather than between populations (Betti Et al. 2009; Betti Et al. 2010; Manica Et al. 2007; Relethford 2001; von Cramon-Taubadel and Lycett 2008; see Figure 14.20). The greatest cranial variation is found among skulls of sub-Saharan African origin, while the least variation is found among populations inhabiting places like Tierra del Fuego at the southern tip of Argentina and Chile. While ancient and historical thinkers previously thought “race” categories could reasonably be determined based on skull dimensions, modern-day analyses using more informative sets of cranial traits simply show that migrations out of Africa and the relative distances between populations can explain a majority of worldwide cranial variation (Betti Et al. 2009).

This same patterning in phenotypic variation has even been found in studies examining shape variation of the pelvis (Betti Et al. 2013; Betti Et al. 2014), the teeth (Rathmann Et al. 2017), and the human bony labyrinth of the ear (Ponce de León Et al. 2018;Figure 14.21). The skeletal morphology of these bones still varies worldwide, but a greater proportion of that variation can still be attributed to the ways in which human populations migrated across the world and exchanged genes with those closer to them rather than those further away. Human skeletal variation in these parts of the body is continuous and nondiscrete. Given the important functions of the cranium and these other skeletal parts, we may infer that the genes that underpin their development have been relatively conserved by neutral evolutionary processes such as genetic drift and gene flow. It is also important to note that while some traits such as height, weight, cranial dimensions, and body composition are determined, in part, by genes, the underlying developmental processes behind these traits are underpinned by complex polygenic mechanisms that have led to the continuous spectrum of variation in such variables among modern-day human populations.
Phenotypic Traits That Reflect Natural Selection
Even though 99.9% of our DNA is the same across all humans worldwide, and many traits reflect neutral processes, there are parts of that remaining 0.1% of the human genome that code for individual and regional differences. Similarly to craniometric analyses that have been conducted in recent decades, human variation in skin colour has also been reassessed using new methods and in light of greater knowledge of biological evolution.
New technologies allow scientists to use colour photometry to sample and quantify the visible wavelength of skin colour, in a way 19th- and 20th-century readers could not. In one report, it was found that 87.9% of global skin colour variation can be attributed to genetic differences between groups, 3.2% to those among local populations within regions, and 8.9% within local populations (Relethford 2002). This apportionment differs significantly and is the reverse situation found in the distribution of genetic differences we see when we examine genetic markers such as blood type–related alleles. However, this pattern of human skin colour worldwide is not surprising, given that we now understand that past selection has occurred for darker skin near the equator and lighter skin at higher latitudes (Jablonski 2004; Jablonski and Chaplin 2000). While most genetic variation reflects neutral variation due to population migrations, geographic isolation, and restricted gene flow dynamics, some human genetic/phenotypic variation is best explained as local adaptation to environmental conditions (i.e., selection). Given that skin colour variation is atypical compared to other genetic markers and biological traits, this, in fact, goes against earlier “race” typologies. This is because recent studies ironically show how so much of genetic variation relates to neutral processes, while skin colour does not. It follows that skin colour cannot be viewed as useful in making inferences about other human traits.

It is also true that some populations have not been studied extensively in skin pigmentation genetics (e.g., African, Austronesian, Melanesian, Southeast Asian, Indigenous American, and Pacific Islander populations, according to Lasisi and Shriver 2018). Earlier dispersals of these populations, and their local genetic varition, will have contributed to worldwide genetic variation, inclusive of skin pigmentation variation. Gene loci we did not previously appreciate as being linked to pigmentation are now being recognized thanks to better tools, more diverse genetic samples, and more accessible datasets (Quillen Et al. 2018). Biological anthropologists look forward to further discoveries elucidating the different selective pressures and population dynamics that influence skin pigmentation evolution.
Social Implications
To finish this chapter, we will consider the social, economic, political, and biological implications of poor understandings of race and the deliberate perpetuation of social and medical racism.
The Black Lives Matter movement (BLM) of 2013 began with the work of racial justice activists and community organizers Alicia Garza, Opal Tometi, and Patrissa Cullors. First incited by the murder of Trayvon Martin, a 17-year-old African American, and the acquittal of the man who shot him, BLM went on to protest against the deaths of numerous Black individuals, most of whom were killed by police officers (for example, Ahmaud Arbery was killed in February of 2020 by two white non-police officers). Some key characteristics of BLM from the start were its decentralized grassroots structure, the role of university students and social media in spreading awareness of the movement, and its embrace of other movements (e.g., climate justice, ending police brutality, feminist campaigns, queer activism, immigration reform, etc.). When George Floyd was murdered by a white police officer on May 25, 2020, the BLM gained new momentum, across 2,000-plus cities in the United States, and among many protesting against historic racism and police brutality in other contexts around the globe. Many in the biological anthropology community have responded to these events with a great dedication to working against systemic racism in society and institutions (American Association of Biological Anthropologists 2020).
BLM continues to be an important movement, as is evidenced in the degree of community organizing, mutual aid efforts, calls for political reform, progress toward curriculum reform and equality, inclusion and diversity (EDI) work in businesses and universities, the removal of monuments honoring historical figures associated with slavery and racism, and many other important actions. Garza (2016) writes: “The reality is that race in the United States operates on a spectrum from black to white … the closer you are to white on that spectrum, the better off you are.” Tometi (2016) has stated: “We need [a human rights movement that challenges systemic racism] because the global reality is that Black people are subject to all sorts of disparities in most of our challenging issues of our day. I think about climate change, and how six of the ten worst impacted nations by climate change are actually on the continent of Africa.” In the words of Cullors (2016), “Black Lives Matter is our call to action. It is a tool to reimagine a world where Black people are free to exist, free to live. It is a tool for our allies to show up differently for us.” We gather from their words the importance of learning from the egregious role that anthropologists have played in the past, recognizing the legacies of “scientific” justifications for eugenics and racism in our society today, and proactively working toward environmental and social equity.
Another major industry that engages in the quantification and interpretation of human variation is medical and clinical work (National Research Council [U.S.] Committee on Human Genome Diversity 1997). Large-scale genomic studies sampling from human populations distributed worldwide have produced detailed knowledge on variation in disease resistance or susceptibility between and within populations. Let’s think about drug companies who develop medicines for Black patients particularly. The predispositions to particular diseases are higher among people of African descent than some pharmaceutical businesses have taken into account. Through targeted sampling of various world groups, clinical geneticists may also identify genetic risk factors of certain common disorders such as chronic heart disease, asthma, diabetes, autoimmune diseases, and behavioural disorders. Having an understanding of population-specific biology is crucial in the development of therapies, medicines, and vaccinations, as not all treatments may be suitable for every human, depending on their genotype. During diagnosis and treatment, it is important to have an evolutionary perspective on gene-environment relationships in patients. Typological concepts of “race” are not useful, given that most racial groups (whether self-identified or not) popularly recognized lack homogeneity and are, in fact, variable. Cystic fibrosis, for instance, occurs in all world populations but can often be underdiagnosed in populations with African ancestry because it is thought of as a “white” disease (Yudell Et al. 2016).
Sociologists, law scholars, and professors of race studies have written extensively on how genetic/technological/medical revolutions impact people of colour. In her book, Fatal Invention: How Science, Politics, and Big Business Re-create Race in the Twenty-First Century (2013), Professor Dorothy E. Roberts writes about how technological advances have been used in resuscitating race as a biological category for dividing humans in essentialist ways (Figure 14.23). She notes how members of law enforcement have engaged in racial profiling, sometimes with the use of machine-learning and facial-recognition technologies. Ancestry-testing services also purport to tell us “what” we are and to insist that this information is “written” in our genes. Such advertising campaigns obscure the nuances of genetic variation with the primary motive of tapping into people’s desire to “know themselves” and driving up profits for their businesses. Commercial genetic testing reinforces the idea that genes map neatly onto race, all while generating massive stores of data in DNA databases. In Roberts’s view, the myth of the biological concept of race being perpetuated in these ways undermines a just society and reproduces racial inequalities.

The COVID-19 pandemic has had a significant impact on the world’s population, particularly people living in the economic Global South and many Black, Indigenous and communities of colour residing in the Global North. We have witnessed disproportionately high numbers of COVID-related deaths and infection cases among marginalized groups. Many immigrants and ethnic minorities in various societies have also experienced scapegoating and blame directed at them for being the source of COVID-19 spread.
To inform us on how to interpret this current worldwide pandemic, historians and anthropologists are looking back at the lessons learned from past instances of racist medicine (discriminatory practices based on broader social discrimination) and medical racism (application of discriminatory practices justified on medical grounds). Historically, who could become doctors and medical professionals was often racialized, gendered, and class specific. This made it difficult for many to overcome prejudices against women, Black people, Indigenous individuals, or other people of colour from becoming doctors and clinical researchers in places such as South Africa and the United States. This, in turn, affects the sorts of information we know about health levels and health outcomes among these very groups. In the past decade, long-overdue attention is finally being paid to how race affects biological outcomes. For instance, researchers have focused on the negative legacies of racial discrimination and racism-induced stress on hormone (im)balances, mental health disorders, cardiovascular disease prevalence, and other health outcomes (Kuzawa and Sweet 2009; Shonkoff, Slopen, and WIlliams 2021; Williams 2018). The technology and standards of protocol in medical testing have been scrutinized (for more on how pulse oximeters were not designed with nonwhite patients in mind, for example, see Sjoding Et al. 2020). Scholars of race and medicine have also written on how illness and disease spread have often been used to perpetuate societal prejudices. This manifests as xenophobic tendencies at a societal level, such as the blaming of “outgroups” and increased “in-group” protectiveness. Overreliance on the idea that people are “inherently” disease carriers due to genetic or biological reasons leads to improper accounting for socioeconomic or infrastructural issues that lead to differential disease prevalence amongst minority communities. (For more on race and COVID, see Tsai 2021 as well as this textbook’s Chapter 16: Contemporary Topics: Human Biology and Health.)
Lastly, consider the changing field of forensic anthropology. In the past, forensic anthropologists ascribed ancestry or racial categories to sets of skeletons, reliant on the belief that different human groups will exhibit biologically “discrete” assortments so as to divide along culturally constructed categories (Sauer 1992). Now, a number of forensic anthropologists have argued that we should abandon these methods, both because it is unscientific and because it further validates and perpetuates this idea that race is biologically meaningful. As scientists, whether we affirm biological race as real has huge influence on the beliefs of members of the public, the judicolegal system, and law enforcement. Not all forensic experts agree with abandoning ancestry estimation. Some prefer to refocus on the neutral or selective causes of human biological variation, and assess how probabilistic it may be to assign bones of certain dimensions to one of several identified racial categories. These debates continue today as this textbook chapter is being written. More details on population affinity may be found in Chapter 17: Forensic Anthropology.
It is important to remember that while it is possible to look for clues about one’s ancestry or geographic origin based on skull morphology, again, the amount of distinctiveness in any given sample makes it impossible to distinguish whether a cranium belongs to one group (Relethford 2009). Individuals can vary in their skeletal dimensions by continental origin, country origin, regional origin, sex, age, environmental factors, and the time period in which they lived, making it difficult to assign individuals to particular categories in a completely meaningful way (Ousley, Jantz, and Freid 2009). When forensic reports and scientific journal articles give an estimation of ancestry, it is crucial to keep in mind that responsible assignments of ancestry will be done through robust statistical testing and stated as a probability estimate. Today, we also live in a more globalized world where a skeletal individual may have been born originally to parents of two separate traditional racial categories. In contexts of great heterogeneity within populations, this definitely adds difficulty to the work of forensic scientists and anthropologists preparing results for the courtroom (genetic testing may be comparatively more helpful in such situations).
Dig Deeper: Measuring FST
Richard Lewontin (1929‒) is a biologist and evolutionary geneticist who authored an article evaluating where the total genetic variation in humans lies. Titled “The Apportionment of Human Diversity” (Lewontin 1972), the article addressed the following question: On average, how genetically similar are two randomly chosen people from the same group when compared to two randomly chosen people from different groups?
Lewontin studied this problem by using genetic data. He obtained data for a large number of different human populations worldwide using 17 genetic markers (including alleles that code for various important enzymes and proteins, such as blood-group proteins). The statistical analysis he ran used a measure of human genetic differences in and among populations known as the fixation index (FST).
Technically, FST can be defined as the proportion of total genetic variance within a subpopulation relative to the total genetic variance from an entire population. Therefore, FST values range from 0 to 1 (or, sometimes you will see this stated as a percentage between 0% and 100%). The closer the FST value of a population (e.g., the world’s population) approaches 1, the higher the degree of genetic differentiation among subpopulations relative to the overall population (see Figure 14.24 for a detailed illustration).

In his article, Lewontin (1972) identified that most of human genetic differences (85.4%) were found within local subpopulations (e.g., the Germans or Easter Islanders), whereas 8.3% were found between populations within continental human groups, and 6.3% were attributable to traditional “race” groups (e.g., “Caucasian” or “Amerind”). These findings have been important for scientifically rejecting the existence of biological races (Long and Kittles 2003).
Talking About Human Biological Variation Going Forward
To conclude, utilizing the term races to describe human biological variation is not accurate or productive. Using a select few hundred genetic loci, or perhaps a number of phenotypic traits, it may be possible to assign individuals to a geographic ancestry, but what constitutes a bounded genetic or geographical grouping is both arbitrary and potentially harmful owing to ethical and historical reasons. The discipline of biological anthropology has moved past typological frameworks that shoehorn continuously variable human populations into discrete and socially constructed subsets. Improvements in the number of markers, the genetic technologies used to study variation, and the number of worldwide populations sampled have led to more nuanced understandings of human variation. It is of utmost importance that scientists make the following points clear to the public:
- Today, we refer to different local human groups as “populations.” What constitutes a population should be carefully defined in scientific reports based on some geographical, linguistic, or cultural criteria and some degree of relativity to other closely or distantly related human groups.
- Humans have significantly less genetic variation than other primates and mammals, and all human beings on Earth share 99.9% of their overall DNA. Some of the remaining 0.1% of human variation varies on a clinal or continuous basis, such as can be seen when looking at ABO blood-type polymorphisms worldwide.
- Many biological characteristics in humans are actually determined nonconcordantly and/or polygenically. Therefore, superiority or inferiority in human behaviour or body form cannot justifiably be linked to fixed and innate differences between groups.
- Genetic distances are correlated with geographic distances among the global human population. This is especially apparent when we consider that genetic variation is highest in sub-Saharan Africa, and average genetic heterogeneity decreases in populations further away from the African continent in accordance with the migratory history of anatomically modern Homo sapiens.
- The effects of gene flow, genetic drift, and population bottlenecking are reflected in some phenotypic traits, such as cranial shape.
- We recognize other traits, like skin colour and lactase persistence, to be the products of many millennia of natural selective pressures influencing human biology from the external environment.
Taken together, genetic analyses of human variation do not support 20th-century (or even-earlier) concepts of race. In discussions about human variation, these genomic results help clarify how biological variation is distributed across the human population today. Taking care to think about and debate the nature of human variation is important, because although the effects and events that produced genetic differences among groups occurred in the ancient past, sociocultural concepts about race and ethnicity continue to have real social, economic, and political consequences.
Beyond talking about variation in the university setting, it is important that teachers, researchers, and students of anthropology recognize and assume the responsibility of influencing public perspectives of human variation. Race-based classification systems were developed during the colonial era, the transatlantic trafficking of kidnapped Africans and the so-called “Scientific Revolution” by the first “anthropologists” and scholars of humankind’s variation. Unfortunately, some of their early ideas have persisted and evolved into present-day lived realities. Some of today’s politicians and socioeconomic bodies have racially charged agendas that promote racism or certain kinds of economic or racial inequalities. As anthropologists, we must acknowledge that while human “races” are not a biological reality, their status as a (misguided) social construction does have real consequences for many people (Antrosio 2011).
In other words, while “race” is a sociocultural invention, the treatment different individuals receive due to their perceived “race” can have significant financial, emotional, sociopolitical, and physiological costs. However—and importantly assuming a “colour-blind” position when it comes to the topics of “race” and ethnicity (especially in political discussions) is actually counterproductive, because the negative social consequences of modern “race” ideas could be ignored, making it harder to examine and address instances of discrimination properly (Wise 2010). Rather than shy away from these topics, we can use our scientific findings to establish socially relevant and biologically accurate ideas concerning human diversity. Today, research into genetic and phenotypic differentiation among and within various human populations continues to expand in its scope, its technological capabilities, its sample sizes, and its ethical concerns. It is thanks to such scientific work done in the past few decades that we now have a deeper understanding not only of how humans vary but also of how we are biologically a rather homogenous, intermixing world population.
Summary
Historically, most concepts of race were shaped by religious and early ‘scientific’ attempts to classify people, many of which justified inequality and racism. Today, biological anthropology emphasizes that human variation is continuous (clinal), polygenic, and shaped by both evolutionary pressures and neutral processes. Misunderstandings of race, however, continue to have serious social and medical consequences; evident in systemic racism and inequities in health care.
Biological anthropologists now play a critical role in discrediting myths about race. By studying human variation, we can begin to understand evolutionary processes, adaptation, and the social implications of differences among human populations. Modern research shows that that humans are far more genetically homogenous than many other species, reinforcing a conclusion that ‘race’ is not a biological reality but a powerful social construct with real effects. Through evolutionary history and genetics, human diversity can be researched and understood, rather than than through racial categories.
Review Questions
- How is the genetic variation of the human species distributed worldwide?
- What evolutionary processes are responsible for producing genotypic/phenotypic variation within and between human populations?
- Should we continue to attribute any value to “race” concepts older than 1950, based on our current understandings of human biological variation?
- How should we communicate scientific findings about human biological variation more accurately and responsibly to those outside the anthropological discipline?
Key Terms
Age of Discovery: A period between the late 1400s and late 1700s when European explorers and ships sailed extensively across the globe in pursuit of new trading routes and territorial conquest.
Ancestry: Biogeographical information about an individual, traced either through the study of an individual’s genome, skeletal characteristics, or some other form of forensic/archaeological evidence. Anthropologists carry out probabilistic estimates of ancestry. They attribute sets of human remains to distinctive “ancestral” groups using careful statistical testing and should report ancestry estimations with statistical probability values.
Binomial nomenclature: A system of naming living things developed by Linnaeus in the 1700s. It employs a scientific name made up of two italicized Latin or Greek words, with the first word capitalized and representative of an organism’s genus and the second word indicating an organism’s species (e.g., Homo sapiens, Australopithecus afarensis, Pongo tapanuliensis, etc.).
Biological anthropology: A branch of study under anthropology (the study of humankind) that focuses on when and where humans and our human ancestors first originated, how we have evolved and adapted globally over time, and the reasons why we see biological variation among humans worldwide today.
Biological determinism: The erroneous concept that an individual’s behavioural characteristics are innate and determined by genes, brain size, or other physiological attributes—and, notably, without the influence of social learning or the environment around the individual during development.
Bony labyrinth: A system of interconnected canals within the auditory (ear- or hearing-related) apparatus, located in the inner ear and responsible for balance and the reception of sound waves.
Cline: A gradient of physiological or morphological change in a single character or allele frequency among a group of species across environmental or geographical lines (e.g., skin colour varies clinally, as, over many generations, human groups living nearer the equator have adapted to have more skin pigmentation).
Continuous variation: This term refers to variation that exists between individuals and cannot be measured using distinct categories. Instead, differences between individuals within a population in relation to one particular trait are measurable along a smooth, continuous gradient.
Cystic fibrosis: A genetic disorder in which one defective gene causes overproduction and buildup of mucus in the lungs and other bodily organs. It is most common in northern Europeans (but also occurs in other world populations).
Ecological niche: The position or status of an organism within its community and/or ecosystem, resulting from the organism’s structural and functional adaptations (e.g., bipedalism, omnivorous diets, lactose digestion, etc.).
Essentialism: A belief or view that an entity, organism, or human grouping has a specific set of characteristics that are fundamentally necessary to its being and classification into definitive categories.
Ethnicity: A term used commonly in an interchangeable way with the term race, complicated because how different people define this term depends on the qualities and characteristics they use to assign a label or identity to themselves and/or others (which may include aspects of family background, skin colour, language(s) spoken, religion, physical proportions, behaviour and temperament, etc.).
Eugenics: A set of beliefs and practices that involves the controlled selective breeding of human populations with the hope of improving their heritable qualities, especially through surgical procedures like sterilization and legal rulings that affect marriage rights for interracial couples.
Gene flow: A neutral (or nonselective) evolutionary process that occurs when genes get shared between populations.
Genetic drift: A neutral evolutionary process in which allele frequencies change from generation to generation due to random chance.
Heterogeneity: The quality of being diverse genetically.
Homogenous: The quality of being uniform genetically.
Human diversity: Human diversity is a measure of variation that may describe how many different forms of human there are, separated or clustered into groups according to some genetic, phenotypic, or cultural trait(s). The term can be applied to culture (in which case humans can be described as significantly diverse) or genetics (in which case humans are not diverse because all humans on Earth share a majority of their genes).
Human variation: Differences in biology, physiology, body chemistry, behaviour, and culture. By measuring these differences, we understand the degrees of variation between individuals, groups, populations, or species.
Isolation-by-distance model: A model that predicts a positive relationship between genetic distances and geographical distances between pairs of populations.
Monogenetic: Pertaining to the idea that the origin of a species is situated in one geographic region or time (as opposed to polygenetic).
Mutation: A gene alteration in the DNA sequence of an organism. As a random, neutral evolutionary process that occurs over the course of meiosis and early cell development, gene mutations are possible sources of variation in any given human gene pool. Genetic mutations that occur in more than 1% of a population are termed polymorphisms.
Natural selection: An evolutionary process whereby certain traits are perpetuated through successive generations, likely owing to the advantages they give organisms in terms of chances of survival and/or reproduction.
Nonconcordance: The fact of genes or traits not varying with one another and instead being inherited independently.
Otherness: In postcolonial anthropology, we now understand “othering” to mean any action by someone or some group that establishes a division between “us” and “them” in relation to other individuals or populations. This could be based on linguistic or cultural differences, and it has largely been based on external characteristics throughout history.
Out-of-Africa model: A model that suggests that all humans originate from one single group of Homo sapiens in (sub-Saharan) Africa who lived between 100,000 and 315,000 years ago and who subsequently diverged and migrated to other regions across the globe.
Physical anthropology: This used to be the more common name given to the subdiscipline of anthropology centred upon the study of human origins, evolution and variation (also see biological anthropology above). This name for the field has gradually become less popular due to two reasons: first, it may not reflect our interests in other aspects of humankind that are not physical (such as those behavioural, cultural and spiritual), and second, using this term popular in the early decades of our field may be viewed by some as harkening back to a time when biological anthropologists conducted their work in unethical ways.
Polygenetic: Having many different ancestries, as in older theories about human origins that involved multiple traditional groupings of humans evolving concurrently in different parts of the world before they merged into one species through interbreeding and/or intergroup warfare. These earlier suggestions have now been overwhelmed by insurmountable evidence for a single origin of the human species in Africa (see the “Out-of-Africa model”).
Polymorphism: A genetic variant within a population (caused either by a single gene or multiple genes) that occurs at a rate of over 1% among the population. Polymorphisms are responsible for variation in phenotypic traits such as blood type and skin colour.
Population: A group of humans living in a particular geographical area, with more local interbreeding within-group than interbreeding with other groups. A limited or restricted amount of gene flow between populations can occur due to geographical, cultural, linguistic, or environmental factors.
Population bottlenecking: An event in which genetic variation is significantly reduced owing to a sharp reduction in population size. This can occur when environmental disaster strikes or as a result of human activities (e.g., genocides or group migrations). An important example of this loss in genetic variation occurred over the first human migrations out of Africa and into other continental regions.
Prejudice: An unjustified attitude toward an individual or group that is not based on reason, whether positive (and showing preference for one group of people over another) or negative (and resulting in harm or injury to others).
Race: The identification of a group based on a perceived distinctiveness that makes that group more similar to each other than they are to others outside the group. This may be based on cultural differences, genetic parentage, physical characteristics, behavioural attributes, or something arbitrarily and socially constructed. As a social or demographic category, perceptions of “race” can have real and serious consequences for different groups of people. This is despite the fact that biological anthropologists and geneticists have demonstrated that all humans are genetically homogenous and that more differences can be found within populations than between them in the overall apportionment of human biological variation. This term is sometimes used interchangeably with ethnicity.
Racism: Any action or belief that discriminates against someone based on perceived differences in race or ethnicity.
Scientific Revolution: A period between the 1400s and 1600s when substantial shifts occurred in the social, technological, and philosophical sense, when a scientific method based on the collection of empirical evidence through experimentation was emphasized and inductive reasoning was used to test hypotheses and interpret their results.
Typological: Of or describing an assortment system that relies on the interpretation of qualitative similarities or differences in the study of variation among objects or people. The categorization of cultures or human groups according to “race” was performed with a typological approach in the earliest practice of anthropology, but this practice has since been discredited and abandoned.
Variance: In statistics, variance measures the dispersal of a set of data around the mean or average value.
For Further Exploration
Videos
American Medical Association (AMA). 2020. “Examining Race-Based Medicine.” YouTube, October 29. Accessed June 4, 2023.
Crenshaw, Kimberlé. 2016. “The Urgency of Intersectionality.” YouTube, December 7. Accessed June 4, 2023.
Golash-Boza, Tanya. 2018. “What Is Race? What Is Ethnicity? Is There a Difference?.” YouTube, October 28. Accessed June 4, 2023.
Lasisi, Tina. 2020. “How Hair Reveals the Futility of Race Categories.” National Museum of Natural History webinar, October 21.
Lasisi, Tina. 2022. “Where Does My Skin Color Come From?.” PBS Terra, August 18. Accessed June 4, 2023.
PBS Origins. 2018. “The Origin of Race in the USA.” YouTube, April 3. Accessed June 4, 2023.
Roberts, Dorothy. 2016. “The Problem with Race-Based Medicine.” YouTube, March 4. Accessed June 4, 2023.
Vox. 2015. “The Myth of Race, Debunked in 3 Minutes.” YouTube, January 13. Accessed June 4, 2023.
Podcast Episodes
Kwong, Emily, and Rebecca Ramirez. 2021. “Here’s a Better Way to Talk about Hair: A 16 Minute Listen with Tina, Lasisi” NPR Short Wave, October 6. Accessed June 4, 2023.
Speaking of Race. 2020. “Race and Health series.” Speaking of Race, April 10. Accessed June 4, 2023.
Websites
Choices Program. 2023. “An Interactive Timeline: Black Activism and the Long Fight for Racial Justice.” Choices Program, Brown University [Interactive Timeline], Updated February, 2023.
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Jonathan M. G. Perry, Ph.D., Western University of Health Sciences
Stephanie L. Canington, Ph.D., University of Pennsylvania
This chapter is a revision from "Chapter 8: Primate Evolution” by Jonathan M. G. Perry and Stephanie L. Canington. In Explorations: An Open Invitation to Biological Anthropology, first edition, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under CC BY-NC 4.0.
Learning Objectives
- Understand the major trends in primate evolution from the origin of primates to the origin of our own species.
- Learn about primate adaptations and how they characterize major primate groups.
- Discuss the kinds of evidence that anthropologists use to find out how extinct primates are related to each other and to living primates.
- Recognize how the changing geography and climate of Earth have influenced where and when primates have thrived or gone extinct.
The first fifty million years of primate evolution was a series of adaptive radiations leading to the diversification of the earliest lemurs, monkeys, and apes. The primate story begins in the canopy and understory of conifer-dominated forests, with our small, furtive ancestors subsisting at night, beneath the notice of day-active dinosaurs.
From the ancient plesiadapiforms (archaic primates) to the earliest groups of true primates (euprimates) (Bloch and Boyer 2002), the origin of our own order is characterized by the struggle for new food sources and microhabitats in the arboreal setting. Climate change forced major extinctions as the northern continents became increasingly dry, cold, and seasonal and as tropical rainforests gave way to deciduous forests, woodlands, and eventually grasslands. Lemurs, lorises, and tarsiers—once diverse groups containing many species—became rare, except for lemurs in Madagascar, where there were no anthropoid competitors and perhaps few predators. Meanwhile, anthropoids (monkeys and apes) likely emerged in Asia and then dispersed across parts of the northern hemisphere, Africa, and ultimately South America. The movement of continents, shifting sea levels, and changing patterns of rainfall and vegetation contributed to the developing landscape of primate biogeography, morphology, and behavior. Today’s primates provide modest reminders of the past diversity and remarkable adaptations of their extinct relatives. This chapter explores the major trends in primate evolution from the origin of the Order Primates to the beginnings of our own lineage, providing a window into these stories from our ancient past.
Major Hypotheses About Primate Origins
For many groups of mammals, there is a key feature that led to their success. A good example is powered flight in bats. Primates lack a feature like this (see Chapter 5). Instead, if there is something unique about primates, it is probably a group of features rather than one single thing. Because of this, anthropologists and paleontologists struggle to describe an ecological scenario that could explain the rise and success of our own order. Three major hypotheses have been advanced to consider the origin of primates and to explain what makes our order distinct among mammals (Figure 9.1); these are described below.

Arboreal Hypothesis
In the 1800s, many anthropologists viewed all animals in relation to humans. That is, animals that were more like humans were considered to be more “advanced” and those lacking humanlike features were considered more “primitive.” This way of thinking was particularly obvious in studies of primates. A more modern way of referring to members of a group that lack certain evolutionary innovations seen in other members is to call them plesiomorphic (literally “anciently shaped”). The state of their morphological features is sometimes referred to as ancestral traits.
Thus, when anthropologists sought features that separate primates from other mammals, they focused on features that were least developed in lemurs and lorises, more developed in monkeys, and most developed in apes (Figure 9.2). Frederic Wood Jones, one of the leading anatomist-anthropologists of the early 1900s, is usually credited with the Arboreal Hypothesis of primate origins (Jones 1916). This hypothesis holds that many of the features of primates evolved to improve locomotion in the trees; this way of getting around is referred to as arboreal. For example, the grasping hands and feet of primates are well suited to gripping tree branches of various sizes and our flexible joints are good for reorienting the extremities in many different ways. A mentor of Jones, Grafton Elliot Smith, had suggested that the reduced olfactory system, acute vision, and forward-facing eyes of primates are adaptations for making accurate leaps and bounds through a complex, three-dimensional canopy (Smith 1912). The forward orientation of the eyes in primates causes the visual fields to overlap, enhancing depth perception, especially at close range. Evidence to support this hypothesis includes the facts that many extant primates are arboreal, and the plesiomorphic members of most primate groups are dedicated arborealists. The Arboreal Hypothesis was well accepted by most anthropologists at the time and for decades afterward.

Visual Predation Hypothesis
In the late 1960s and early 1970s, Matt Cartmill studied and tested the idea that the characteristic features of primates evolved in the context of arboreal locomotion. Cartmill noted that squirrels climb trees (and even vertical walls) very effectively, even though they lack some of the key adaptations of primates. As members of the Order Rodentia, squirrels also lack the hand and foot anatomy of primates. They have claws instead of flattened nails and their eyes face more laterally than those of primates. Cartmill reasoned that there must be some other explanation for the unique traits of primates. He noted that some nonarboreal animals share at least some of these traits with primates; for example, cats and predatory birds have forward-facing eyes that enable visual field overlap. Cartmill suggested that the unique suite of features in primates is an adaptation to detecting insect prey and guiding the hands (or feet) to catch insects (Cartmill 1972). His hypothesis emphasizes the primary role of vision in prey detection and capture; it is explicitly comparative, relying on form-function relationships in other mammals and nonmammalian vertebrates. According to Cartmill, many of the key features of primates evolved for preying on insects in this special manner (Cartmill 1974).
Angiosperm-Primate Coevolution Hypothesis
The visual predation hypothesis was unpopular with some anthropologists. One reason for this is that many primates today are not especially predatory. Another is that, whereas primates do seem well adapted to moving around in the smallest, terminal branches of trees, insects are not necessarily easier to find there. A counterargument to the visual predation hypothesis is the angiosperm-primate coevolution hypothesis. Primate ecologist Robert Sussman (Sussman 1991) argued that the few primates that eat mostly insects often catch their prey on the ground rather than in tree branches. Furthermore, predatory primates often use their ears more than their eyes to detect prey. Finally, most early primate fossils show signs of having been omnivorous rather than insectivorous. Instead, he argued, the earliest primates were probably seeking fruit. Fruit (and flowers) of angiosperms (flowering plants) often develop in the terminal branches. Therefore, any mammal trying to access those fruits must possess anatomical traits that allow them to maintain their hold on thin branches and avoid falling while reaching for the fruits. Primates likely evolved their distinctive visual traits and extremities in the Paleocene (approximately 65 million to 54 million years ago) and Eocene (approximately 54 million to 34 million years ago) epochs, just when angiosperms were going through a revolution of their own—the evolution of large, fleshy fruit that would have been attractive to a small arboreal mammal. Sussman argued that, just as primates were evolving anatomical traits that made them more efficient fruit foragers, angiosperms were also evolving fruit that would be more attractive to primates to promote better seed dispersal. This mutually beneficial relationship between the angiosperms and the primates was termed coevolution or more specifically diffuse coevolution.
At about the same time, D. Tab Rasmussen noted several parallel traits in primates and the South American woolly opossum, Caluromys. He argued that early primates were probably foraging on both fruits and insects (Rasmussen 1990). As is true of Caluromys today, early primates probably foraged for fruits in the terminal branches of angiosperms, and they probably used their visual sense to aid in catching insects. Insects are also attracted to fruit (and flowers), so these insects represent a convenient opportunity for a primarily fruit-eating primate to gather protein. This solution is a compromise between the visual predation hypothesis and the angiosperm-primate coevolution hypothesis. It is worth noting that other models of primate origins have been proposed, and these include the possibility that no single ecological scenario can account for the origin of primates.
The Origins of Primates
Paleocene: Mammals in the Wake of Dinosaur Extinctions
Placental mammals, including primates, originated in the Mesozoic Era (approximately 251 million to 65.5 million years ago), the Age of Dinosaurs. During this time, most placental mammals were small, probably nocturnal, and probably avoided predators via camouflage and slow, quiet movement. It has been suggested that the success and diversity of the dinosaurs constituted a kind of ecological barrier to Mesozoic mammals. The extinction of the dinosaurs (and many other organisms) at the end of the Cretaceous Period (approximately 145.5–65.5 million years ago) might have opened up these ecological niches, leading to the increased diversity and disparity in mammals of the Tertiary Period (approximately 65.5–2.5 million years ago).
The Paleocene was the first epoch in the Age of Mammals. Soon after the Cretaceous-Tertiary (K-T) extinction event, new groups of placental mammals appear in the fossil record. Many of these groups achieved a broad range of sizes and lifestyles as well as a great number of species before declining sometime in the Eocene (or soon thereafter). These groups were ultimately replaced by the modern orders of placental mammals (Figure 9.3). It is unknown whether these replacements occurred gradually, for example by competitive exclusion, or rapidly, perhaps by sudden geographic dispersals with replacement. In some senses, the Paleocene might have been a time of recovery from the extinction event; it was cooler and more seasonal globally than the subsequent Eocene.

Plesiadapiforms, the Archaic Primates
The Paleocene epoch saw the emergence of several families of mammals that have been implicated in the origin of primates. These are the plesiadapiforms, which are archaic primates, meaning they possessed some primate features and lacked others. The word plesiadapiform means “almost adapiform,” a reference to some similarities between some plesiadapiforms and some adapiforms (or adapoids; later-appearing true primates)—mainly in the molar teeth. Because enamel fossilizes better than other parts of the body, the molar teeth are the parts most often found and first discovered for any new species. Thus, dental similarities were often the first to be noticed by early mammalian paleontologists, partly explaining why plesiadapiforms were thought to be primates. Major morphological differences between plesidapiforms and euprimates (true primates) were observed later when more parts of plesiadapiform skeletons were discovered. Many plesiadapiforms have unusual anterior teeth and most have digits possessing claws rather than nails. So far, no plesiadapiform ever discovered has a postorbital bar (seen in extant strepsirrhines) or septum (as seen in haplorhines), and whether or not the auditory bulla was formed by the petrosal bone remains unclear for many plesiadapiform specimens. Nevertheless, there are compelling reasons (partly from new skeletal material) for including plesiadapiforms within the Order Primates.
Geographic and Temporal Distribution
Purgatorius is generally considered to be the earliest primate. This Paleocene mammal is known from teeth that are very plesiomorphic for a primate. It has some characteristics that suggest it is a basal plesiadapiform, but there is very little to link it specifically with euprimates (see Clemens 2004). Its ankle bones suggest a high degree of mobility, signaling an arboreal lifestyle (Chester et al. 2015). Purgatorius is plesiomorphic enough to have given rise to all primates, including the plesiadapiforms. However, new finds suggest that this genus was more diverse and had more differing tooth morphologies than previously appreciated (Wilson Mantilla et al. 2021). Plesiadapiform families were numerous and diverse during parts of the Paleocene in western North America and western Europe, with some genera (e.g., Plesiadapis; see Figure 9.4) living on both continents (Figure 9.5). Thus, there were probably corridors for plesiadapiform dispersal between the two continents, and it stands to reason that these mammals were living all across North America, including in the eastern half of the continent and at high latitudes. A few plesiadapiforms have been described from Asia (e.g., Carpocristes), but the affinities of these remain uncertain.
|
Family
|
Genera
|
Morphology
|
Location
|
Age1
|
|
Paromomyidae |
Ignacius |
Long, dagger-like, lower incisor. |
North America and Europe |
Early Paleocene to Late Eocene |
|
Carpolestidae |
Carpolestes |
Plagiaulacoid dentition. Limb adaptations to terminal branch feeding. Grasping big toe. |
North America, Europe, and Asia |
Middle Paleocene to Early Eocene |
|
Plesiadapidae |
Plesiadapis |
Mitten-like upper incisor. Diastema. Large body size for group. |
North America and Europe |
Middle Paleocene to Early Eocene |
|
1 Derived from Fleagle 2013. |
|
|||

General Morphological Features
Although there is much morphological variation among the families of plesiadapiforms, some common features unite the group. Most plesiadapiforms were small, the largest being about three kilograms (approximately 7 lbs.; Plesiadapis cookei). They had small brains and fairly large snouts, with eyes that faced more laterally than in euprimates. Many species show reduction and/or loss of the canine and anterior premolars, with the resulting formation of a rodent-like diastema (a pronounced gap between the premolars and the incisors, with loss of at least the canine); this probably implies a herbivorous diet. Some families appear to have had very specialized diets, as suggested by unusual tooth and jaw shapes.
Arguably the most interesting and unusual family of plesiadapiforms is the Carpolestidae. They are almost exclusively from North America (with a couple of possible members from Asia), and mainly from the Middle and Late Paleocene. Their molars are not very remarkable, being quite similar to those of some other plesiadapiforms (e.g., Plesiadapidae). However, their lower posterior premolars (p4) are laterally compressed and blade-like with vertical serrations topped by tiny cuspules. This unusual dental morphology is termed plagiaulacoid (Simpson 1933). The upper premolar occlusal surfaces are broad and are covered with many small cuspules; the blade-like lower premolar might have cut across these cuspules, between them, or both.

Many plesiadapiforms have robust limb bones with hallmarks of arboreality. Instead of having nails, most taxa had sharp claws on most or all of the digits. The extremities show grasping abilities comparable to those of primates and some arboreal marsupials. Nearly complete skeletons have yielded a tremendous wealth of information on locomotor and foraging habits. Many plesiadapiforms appear to have been able to cling to vertical substrates (like a broad tree trunk) using their sharp claws, propelling themselves upward using powerful hindlimbs, bounding along horizontal supports, grasping smaller branches, and moving head-first down tree trunks. In carpolestids in particular, the skeleton appears to have been especially well adapted to moving slowly and carefully in small terminal branches (Figure 9.6).
Dig Deeper: Debate: Relationship of Plesiadapiforms to True Primates
In the middle of the twentieth century, treeshrews (Order Scandentia) were often considered part of the Order Primates, based on anatomical similarities between some treeshrews and primates. For many people, plesiadapiforms represented intermediates between primates and treeshrews, so plesiadapiforms were included in Primates as well.
Studies of reproduction and brain anatomy in treeshrews and lemurs suggested that treeshrews are not primates (e.g., Martin 1968). This was soon followed by the suggestion to also expel plesiadapiforms (Martin 1972) from the Order Primates. Like treeshrews, plesiadapiforms lack a postorbital bar, nails, and details of the ear region that characterize true primates. Many paleoanthropologists were reluctant to accept this move to banish plesiadapiforms (e.g., F. S. Szalay, P. D. Gingerich).
Later, K. Christopher Beard (1990) found that in some ways, the digits of paromomyid plesiadapiforms are actually more similar to those of dermopterans (Order Dermoptera), the closest living relatives of primates, than they are to those of primates themselves (but see Krause 1991). At the same time, Richard Kay and colleagues (1990) found that cranial circulation patterns and auditory bulla morphology in the paromomyid, Ignacius (see Figure 9.4), are more like those of dermopterans than of primates.
For many anthropologists, this one-two punch effectively removed plesiadapiforms from the Order Primates. In the last two decades, the tide of opinion has turned again, with many researchers reinstating plesiadapiforms as members of the Order Primates. New and more complete specimens demonstrate that the postcranial skeletons of plesiadapiforms, including the hands and feet, were primate-like, not dermorpteran-like (Bloch and Boyer 2002, 2007). New fine-grained CT scans of relatively complete plesiadapiform skulls revealed that they share some key traits with primates to the exclusion of other placental mammals (Bloch and Silcox 2006). Most significant was the suggestion that Carpolestes simpsoni possessed an auditory bulla formed by the petrosal bone, like in all living primates.
The debate about the status of plesiadapiforms continues, owing to a persistent lack of key bones in some species and owing to genuine complexity of the anatomical traits involved. Maybe plesiadapiforms were the ancestral stock from which all primates arose, with some plesiadapiforms (e.g., carpolestids) nearer to the primate stem than others.
Adapoids and Omomyoids, the First True Primates
Geographic and Temporal Distribution
The first universally accepted fossil primates are the adapoids (Superfamily Adapoidea) and the omomyoids (Superfamily Omomyoidea). These groups become quite distinct over evolutionary time, filling mutually exclusive niches for the most part. However, the earliest adapoids are very similar to the earliest omomyoids.
The adapoids were mainly diurnal and herbivorous, with some achieving larger sizes than any plesiadapiforms (10 kg; 22 lbs.). By contrast, the omomyoids were mainly nocturnal, insectivorous and frugivorous, and small.
Both groups appear suddenly at the start of the Eocene, where they are present in western North America, western Europe, and India (Figure 9.7). This wide dispersal of early primates was probably due to the presence of rainforest corridors extending far into northern latitudes.

In North America and Europe, both groups achieved considerable diversity in the Middle Eocene, then mostly died out at the end of that epoch (Figure 9.8). In some Eocene rock formations in the western United States, adapoids and omomyoids make up a major part of the mammalian fauna. The Eocene of India has yielded a modest diversity of euprimates, some of which are so plesiomorphic that it is difficult to know whether they are adapoids or omomyoids (or even early anthropoids).
|
Family
|
Genera
|
Morphology
|
Location
|
Age1
|
|
Cercamoniidae |
Donrussellia |
Variable in tooth number and jaw shape. |
Europe and Asia |
Early to Late Eocene |
|
Asiadapidae2 |
Asiadapis |
Plesiomorphic teeth and jaw resemble early Omomyids. |
Asia |
Early Eocene |
|
Caenopithecidae3 |
Darwinius |
Robust jaws with crested molars. Fewer premolars. |
Europe, Africa, North America, and Asia |
Middle to Late Eocene |
|
Adapidae |
Adapis |
Fused mandible. Long molar crests. Large size and large chewing muscles. |
Europe |
Late Eocene to Early Oligocene |
|
Sivaladapidae |
Sivaladapis |
Some large with robust jaws. |
Asia |
Middle Eocene to Late Miocene |
|
Notharctidae4 |
Notharctus |
Canine sexual dimorphism. Lemur-like skull. Clinging and leaping adaptations. |
North America and Europe |
Early to Middle Eocene |
|
Omomyidae5 |
Teilhardina |
Small, nocturnal, frugivorous or insectivorous. Tarsier-like skull in some. |
North America, Europe, and Asia |
Early Eocene to Early Miocene |
|
Microchoeridae6 |
Necrolemur |
Long bony ear tubes. Tarsier-like lower limb adaptations for leaping. |
Europe and Asia |
Early Eocene to Early Oligocene |
|
1 Derived from Fleagle 2013. 2 See Dunn et al. 2016 and Rose et al. 2018. 3 See Kirk and Williams 2011 and Seiffert et al. 2009. 4 See Gregory 1920. 5 See Beard and MacPhee 1994 and Strait 2001. 6 See Schmid 1979. |
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Adapoids and omomyoids barely survived the Eocene-Oligocene extinctions, when colder temperatures, increased seasonality, and the retreat of rainforests to lower latitudes led to changes in mammalian biogeography. In North America, one genus (originally considered an omomyoid but recently reclassified as Adapoidea) persisted until the Miocene: Ekgmowechashala (Rose and Rensberger 1983). This taxon has highly unusual teeth and might have been a late immigrant to North America from Asia. In Asia, one family of adapoids, the Sivaladapidae, retained considerable diversity as late as the Late Miocene.
Adapoid Diversity
Adapoids were very diverse, particularly in the Eocene of North America and Europe. They can be divided into six families, with a few species of uncertain familial relationship. As a group, adapoids have some features in common, although much of what they share is plesiomorphic. Important features include the hallmarks of euprimates: postorbital bar, flattened nails, grasping extremities, and a petrosal bulla (Figures 9.9 and 9.10). In addition, some adapoids retain the ancestral dental formula of 2.1.4.3; that is, in each quadrant of the mouth, there are two incisors, one canine, four premolars, and three molars. In general, the incisors are small compared to the molars, but the canines are relatively large, with sexual dimorphism in some species. Cutting crests on the molars are well developed in some species, and the two halves of the mandible were fused at the midline in some species. Some adapoids were quite small (Anchomomys at a little over 100 g), and some were quite large (Magnadapis at 10 kg; 22 lbs.). Furthermore, the spaces and attachment features for the chewing muscles were truly enormous in some species, suggesting that these muscles were very large and powerful. Taken together, this suggests an overall adaptive profile of diurnal herbivory. The canine sexual dimorphism in some species suggests a possible mating pattern of polygyny, as males in polygynous primate species often compete with each other for mates and have especially large canine teeth.


Omomyoid Diversity
Like adapoids, omomyoids appeared suddenly at the start of the Eocene and then became very diverse with most species dying out before the Oligocene. Omomyoids are known from thousands of jaws with teeth, relatively complete skulls for about a half-dozen species, and very little postcranial material. Omomyoids were relatively small primates, with the largest being less than three kilograms (approximately 7 lbs.; Macrotarsius montanus). All known crania possess a postorbital bar, which in some has been described as “incipient closure.” Some—but not all—known crania have an elongated bony ear tube extending lateral to the location of the eardrum, a feature seen in living tarsiers and catarrhines. The anterior teeth tend to be large, with canines that are usually not much larger than the incisors. Often it is difficult to distinguish closely related species using molar morphology, but the premolars tend to be distinct from one species to another. The postcranial skeleton of most omomyoids shows hallmarks of leaping behavior reminiscent of that of tarsiers. In North America, omomyoids became very diverse and abundant. In fact, omomyoids from Wyoming are sufficiently abundant and from such stratigraphically controlled conditions that they have served as strong evidence for the gradual evolution of anatomical traits over time (Rose and Bown 1984).
Teilhardina (Figure 9.11; see Figure 9.2) is one of the earliest and arguably the most plesiomorphic of omomyoids. Teilhardina has several species, most of which are from North America, with one from Europe (T. belgica) and one from Asia (T. asiatica). The species of this genus are anatomically similar and the deposits from which they are derived are roughly contemporaneous. Thus, this small primate likely dispersed across the northern continents very rapidly (Smith et al. 2006).

The Emergence of Modern Primate Groups
Origins of Crown Strepsirrhines
Until the turn of this century, very little was known about the origins of the crown (living) strepsirrhines. The Quaternary record of Madagascar contains many amazing forms of lemurs, including giant sloth-like lemurs, lemurs with perhaps monkey-like habits, lemurs with koala-like habits, and even a giant aye-aye (Godfrey and Jungers 2002). However, in Madagascar, early Tertiary continental sediments are lacking, and there is no record of lemur fossils before the Pleistocene.
The fossil record of galagos is slightly more informative. Namely, there are Miocene African fossils that are very likely progenitors of lorisids (Simpson 1967). However, these are much like modern galagos and do not reveal anything about the relationship between crown strepsirrhines and Eocene fossil primates (but see below regarding Propotto). A similar situation exists for lorises in Asia: there are Miocene representatives, but these are substantially like modern lorises. The discovery of the first definite toothcomb canine (a hallmark of stresirrhines) in 2003 provided the “smoking gun” for the origin of crown strepsirrhines (Seiffert et al. 2003). Recently, several other African primates have been recognized as having strepsirrhine affinities (Marivaux et al. 2013; Seiffert 2012). The enigmatic Fayum primate Plesiopithecus is known from a skull that has been compared to aye-ayes and to lorises (Godinot 2006; Simons and Rasmussen 1994a).
The now-recognized diversity of stem strepsirrhines from the Eocene and Oligocene of Afro-Arabia is strong evidence to suggest that strepsirrhines originated in that geographic area. This implies that lorises dispersed to Asia subsequent to an African origin. It is unknown what the first strepsirrhines in Madagascar were like. However, it seems likely that the lemuriform-lorisiform split occurred in continental Africa, followed by dispersal of lemuriform stock to Madagascar. Recent evidence suggests that Propotto, a Miocene primate from Kenya originally described as a potto antecedent, actually forms a clade with Plesiopithecus and the aye-aye; this might suggest that strepsirrhines dispersed to Madagascar from continental Africa more than once (Gunnell et al. 2018).
The Fossil Record of Tarsiers
Tarsiers are so unusual that they fuel major debates about primate taxonomy. Tarsiers today are moderately diverse but geographically limited and not very different in their ecological habits—especially considering that the split between them and their nearest living relative probably occurred over 50 million years ago. If omomyoids are excluded, then the fossil record of tarsiers is very limited. Two fossil species from the Miocene of Thailand have been placed in the genus Tarsius, as has an Eocene fossil from China (Beard et al. 1994). These, and Xanthorhysis from the Eocene of China, are all very tarsier-like. In fact, it is striking that Tarsius eocaenus from China was already so tarsier-like as early as the Eocene. This suggests that tarsiers achieved their current morphology very early in their evolution and have remained more or less the same while other primates changed dramatically. Two additional genera, Afrotarsius from the Oligocene of Egypt and Libya and Afrasia from the Eocene of Myanmar, have also been implicated in tarsier origins, though the relationship between them and tarsiers is unclear (Chaimanee et al. 2012). More recently, a partial skeleton of a small Eocene primate from China, Archicebus achilles (dated to approximately 55.8 million to 54.8 million years ago), was described as the most basal tarsiiform (Ni et al. 2013). This primate is reconstructed as a diurnal insectivore and an arboreal quadruped that did some leaping—but not to the specialized degree seen in living tarsiers. The anatomy of the eye in living tarsiers suggests that their lineage passed through a diurnal stage, so Archicebus (and diurnal omomyoids) might represent such a stage.
Climate Change and the Paleogeography of Modern Primate Origins
Changing global climate has had profound effects on primate dispersal patterns and ecological habits over evolutionary time. Primates today are strongly tied to patches of trees and particular plant parts such as fruits, seeds, and immature leaves. It is no surprise, then, that the distribution of primates mirrors the distribution of forests. Today, primates are most diverse in the tropics, especially in tropical rainforests. Global temperature trends across the Tertiary have affected primate ranges. Following the Cretaceous-Tertiary extinction event, cooler temperatures and greater seasonality characterized the Paleocene. In the Eocene, temperatures (and probably rainfall) increased globally and rainforests likely extended to very high latitudes. During this time, euprimates became diverse. With cooling and increased aridity at the end of the Eocene, many primate extinctions occurred in the northern continents and the surviving primates were confined to lower latitudes in South America, Afro-Arabia, Asia, and southern Europe. Among these survivors are the progenitors of the living groups of primates: lemurs and lorises, tarsiers, platyrrhines (monkeys of the Americas), and catarrhines (monkeys and apes of Africa and Asia) (Figure 9.12).

Competing Hypotheses for the Origin of Anthropoids
There is considerable debate among paleoanthropologists as to the geographic origins of anthropoids. In addition, there is debate regarding the source group for anthropoids. Three different hypotheses have been articulated in the literature. These are the adapoid origin hypothesis, the omomyoid origin hypothesis, and the tarsier origin hypothesis (Figure 9.13).

Adapoid Origin Hypothesis
Resemblances between some adapoids and some extant anthropoids include fusion of the mandibular symphysis, overall robusticity of the chewing system, overall large body size, features that signal a diurnal lifestyle (like relatively small eye sockets), and ankle bone morphology. Another feature in common is canine sexual dimorphism, which is present in some species of adapoids (probably) and in several species of anthropoids.
These features led some paleoanthropologists in the last half of the 20th century to suggest that anthropoids came from adapoid stock (Gingerich 1980; Simons and Rasmussen 1994b). One of the earliest supporters of the link between adapoids and anthropoids was Hans Georg Stehlin, who described much of the best material of adapoids and compared these Eocene primates to South American monkeys (Stehlin 1912). In more recent times, the adapoid origin hypothesis was reinforced by resemblances between these European adapoids (especially Adapis and Leptadapis) and some early anthropoids from the Fayum Basin (e.g., Aegyptopithecus, see below; Figure 9.14).
|
Family
|
Genera
|
Morphology
|
Location
|
Age1
|
|
Propliopithecidae2 |
Aegyptopithecus |
Large size. Cranial sexual dimorphism, large canines. Robust jaws and rounded molars. Partially ossified ear tube (in some). Robust skeleton; quadruped. |
Africa |
Late Eocene to Early Oligocene |
|
Parapithecidae3 |
Apidium |
Medium size. Retention of three premolars per quadrant. Rounded molars and premolars with large central cusps. Adaptations for leaping in the lower limb. |
Africa |
Late Eocene to Late Oligocene |
|
Proteopithecidae4 |
Proteopithecus |
Small size. Retention of three premolars per quadrant. Arboreal quadrupeds that ate fruit. |
Africa |
Late Eocene |
|
Oligopithecidae5 |
Catopithecus |
Small size. Skull has postorbital septum and unfused mandible. Deep jaws. Diet of fruits. Generalized quadruped. |
Africa |
Late Eocene |
|
Eosimiidae |
Eosimias |
Deep jaw with vertical unfused symphysis. Pointed incisors and canines. Crowded premolars. |
Asia |
Middle Eocene |
|
Amphipithecidae6 |
Pondaungia |
Deep jaws. Molars generally rounded with wide basins. |
Asia |
Middle Eocene to Early Oligocene |
|
1 Derived from Fleagle 2013. 2 See Gebo and Simons 1987 and Simons et al. 2007. 3 See Feagle and Simons 1995 and Simons 2001. 4 See Simons and Seiffert 1999. 5 See Simons and Rasmussen 1996. 6 See Kay et al. 2004. |
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Unfortunately for the adapoid hypothesis, most of the shared features listed above probably emerged independently in the two groups as adaptations to a diet of hard and/or tough foods. For example, fusion of the mandibular symphysis likely evolved as a means to strengthen the jaw against forces that would pull the two halves away from each other, in the context of active chewing muscles on both sides of the head generating great bite forces. This context would also favor the development of robust jaws, large chewing muscles, shorter faces, and some other features shared by some adapoids and some anthropoids.
As older and more plesiomorphic anthropoids were found in the Fayum Basin, it became clear that the earliest anthropoids from Africa did not possess these features of jaw robusticity (Seiffert et al. 2009). Furthermore, many adapoids never evolved these features. Fusion of the mandibular symphysis in adapoids is actually quite different from that in anthropoids and probably occurred during juvenile development in the former (Beecher 1983; Ravosa 1996). Eventually, the adapoid origin hypothesis fell out of favor among most paleoanthropologists, although the description of Darwinius is a recent revival of that idea (Franzen et al. 2009; but see Seiffert et al. 2009, Williams et al. 2010b).
Omomyoid Origin Hypothesis
Similarities in cranial and hindlimb morphology between some omomyoids and extant tarsiers have led to the suggestion that tarsiers arose from some kind of omomyoid. In particular, Necrolemur has many features in common with tarsiers, as does the North American Shoshonius, which is known from a few beautifully preserved (although distorted) crania. Tarsiers and Shoshonius share exclusively some features of the base of the cranium; however, Shoshonius does not have any sign of postorbital closure, and it lacks the bony ear tube of tarsiers. Nevertheless, some of the resemblances between some omomyoids and tarsiers suggest that tarsiers might have originated from within the Omomyoidea (Beard 2002; Beard and MacPhee 1994). In this scenario, although living tarsiers and living anthropoids might be sister taxa, they might have evolved from different omomyoids, possibly separated from each other by more than 50 million years of evolution, or from anthropoids evolved from some non-omomyoid fossil group. The arguments against the omomyoid origin hypothesis are essentially the arguments for the tarsier origin hypothesis (see below). Namely, tarsiers and anthropoids share many features (especially of the soft tissues) that must have been retained for many millions of years or must have evolved convergently in the two groups. Furthermore, a key hard-tissue feature shared between the two extant groups, the postorbital septum, was not present in any omomyoid. Therefore, that feature must have arisen convergently in the two extant groups or must have been lost in omomyoids. Neither scenario is very appealing, although recent arguments for convergent evolution of the postorbital septum in tarsiers and anthropoids have arisen from embryology and histology of the structure (DeLeon et al. 2016).
Tarsier Origin Hypothesis
Several paleoanthropologists have suggested that there is a relationship between tarsiers and anthropoids to the exclusion of omomyoids and adapoids (e.g., Cartmill and Kay 1978; Ross 2000; Williams and Kay 1995). Tarsiers and anthropoids today share several traits, including many soft-tissue features related to the olfactory system (e.g., the loss of a hairless external nose and loss of the median cleft running from the nose to the mouth, as possessed by strepsirrhines), and aspects of the visual system (e.g., the loss of a reflective layer at the back of the eye, similarities in carotid circulation to the brain, and mode of placentation). Unfortunately, none of these can be assessed directly in fossils. Some bony similarities between tarsiers and anthropoids include an extra air-filled chamber below the middle ear cavity, reduced bones within the nasal cavity, and substantial postorbital closure; these can be assessed in fossils, but the distribution of these traits in omomyoids does not yield clear answers. Furthermore, several similarities between tarsiers and anthropoids are probably due to similarities in sensory systems, which might have evolved in parallel for ecological reasons. Although early attempts to resolve the crown primates with molecular data were sometimes equivocal or in disagreement with one another, more recent analyses (including those of short interspersed elements) suggest that tarsiers and anthropoids are sister groups to the exclusion of lemurs and lorises (Williams et al. 2010a). However, this does not address omomyoids, all of which are far too ancient for DNA extraction.
The above three hypotheses are not the only possibilities for anthropoid origins. It may be that anthropoids are neither the closest sister group of tarsiers, nor evolved from adapoids or omomyoids. In recent years, two new groups of Eocene Asian primates have been implicated in the origin of anthropoids: the eosimiids and the amphipithecids. It is possible that one or the other of these two groups gave rise to anthropoids. Regardless of the true configuration of the tree for crown primates, the three major extant groups probably diverged from each other quite long ago (Seiffert et al. 2004).
Early Anthropoid Fossils in Africa


The classic localities yielding the greatest wealth of early anthropoid fossils are those from the Fayum Basin in Egypt (Simons 2008; Figure 9.15). The Fayum is a veritable oasis of fossil primates in an otherwise spotty early Tertiary African record. Since the 1960s, teams led by E. L. Simons have discovered several new species of early anthropoids, some of which are known from many parts of the skeleton and several individuals (Figure 9.16).
The Fayum Jebel Qatrani Formation and Birket Qarun Formation between them have yielded a remarkable array of terrestrial, arboreal, and aquatic mammals. These include ungulates, bats, sea cows, elephants, hyraces, rodents, whales, and primates. Also, many other vertebrates, like water birds, were present. The area at the time of deposition (Late Eocene through Early Oligocene) was probably very wet, with slow-moving rivers, standing water, swampy conditions, and lots of trees (see Bown and Kraus 1988). In short, it was an excellent place for primates.
General Morphology of Anthropoids
The anthropoids known from the Fayum (and their close relatives from elsewhere in East Africa and Afro-Arabia) bear many of the anatomical hallmarks of extant anthropoids; however, there are plesiomorphic forms in several families that lack one or more anthropoid traits. All Fayum anthropoids known from skulls possess postorbital closure, most had fused mandibular symphyses, and most had ring-like ectotympanic bones. Tooth formulae were generally either 2.1.3.3 or 2.1.2.3. Fayum anthropoids ranged in size from the very small Qatrania and Biretia (less than 500 g) to the much-larger Aegyptopithecus (approximately 7 kg; 15 lbs.). Fruit was probably the main component of the diet for most or all of the anthropoids, with some of them supplementing with leaves (Kay and Simons 1980; Kirk and Simons 2001; Teaford et al. 1996). Most Fayum anthropoids were probably diurnal above-branch quadrupeds. Some of them (e.g., Apidium; see Figure 9.14) were probably very good leapers (Gebo and Simons 1987), but none show specializations for gibbon-style suspensory locomotion. Some of the Fayum anthropoids are known from hundreds of individuals, permitting the assessment of individual variation, sexual dimorphism, and in some cases growth and development. The description that follows provides greater detail for the two best known Fayum anthropoid families, the Propliopithecidae and the Parapithecidae; the additional families are summarized briefly.
Fayum Anthropoid Families
The Propliopithecidae (see Figure 9.14) include the largest anthropoids from the fauna, and they are known from several crania and some postcranial elements. They have been suggested to be stem catarrhines, although perhaps near the split between catarrhines and platyrrhines. The best known propliopithecid is Aegyptopithecus, known from many teeth, crania, and postcranial elements (Figure 9.17) .

Parapithecidae are an extremely abundant and unusual family of anthropoids from the Fayum. The parapithecid Apidium is known from many jaws with teeth, crushed and distorted crania, and several skeletal elements. Parapithecus is known from cranial material including a beautiful, undistorted cranium. This genus shows extreme reduction of the incisors, including complete absence of the lower incisors in P. grangeri (Simons 2001). This trait is unique among primates. Parapithecids were once thought to be the ancestral stock of platyrrhines; however, their platyrrhine-like features are probably ancestral retentions, so the most conservative approach is to consider them stem anthropoids.
The Proteopithecidae were small frugivores that probably mainly walked along horizontal branches on all fours. They are considered stem anthropoids. The best known genus, Proteopithecus, is represented by dentitions, crania, and postcranial elements.
The Oligopithecidae share a mixture of traits that makes them difficult to classify more specifically within anthropoids. The best known member, Catopithecus, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion. They share the catarrhine tooth formula of 2.1.2.3 and have a canine honing complex that involves the anterior lower premolar. The postcranial elements known for the group suggest generalized arboreal quadrupedalism. The best known member, Catopithecus, is known from crania that demonstrate a postorbital septum and from mandibles that lack symphyseal fusion (Simons and Rasmussen 1996). The jaws are deep, with broad muscle attachment areas and crested teeth. Catopithecus was probably a little less than a kilogram in weight.
Other genera of putative anthropoids from the Fayum include the very poorly known Arsinoea, the contentious Afrotarsius, and the enigmatic Nosmips. The last of these possesses traits of several major primate clades and defies classification (Seiffert et al. 2010).
Early Anthropoid Fossils in Asia
For the last half of the 1900s, researchers believed that Africa was the unquestioned homeland of early anthropoids (see Fleagle and Kay 1994). However, two very different groups of primates from Asia soon began to change that. One was an entirely new discovery (Eosimiidae), and the other was a poorly known group discovered decades prior (Amphipithecidae). Soon, attention on anthropoid origins began to shift eastward (see Ross and Kay 2004; Simons 2004). If anthropoids arose in Asia instead of Africa, then this implies that the African early anthropoids either emigrated from Asia or evolved their anthropoid traits in parallel with living anthropoids.
Eosimiids
First described in the 1990s, the eosimiids are best represented by Eosimias (see Figure 9.14; Figure 9.18). This tiny “dawn monkey” is known from relatively complete jaws with teeth, a few small fragments of the face, and some postcranial elements (Beard et al. 1994; Beard et al. 1996; Gebo et al. 2000). Eosimias (along with the other less-well-known genera in its family) bears some resemblance to tarsiers as well as anthropoids. Unfortunately, no good crania are known for this family, and the anatomy of, for example, the posterior orbital margin could be very revealing as to higher-level relationships.

Amphipithecids
Amphipithecids are small- to medium-size primates (up to 10 kg; 22 lbs.). Most are from the Eocene Pondaung Formation in Myanmar (Early–Middle Eocene), but one genus is known from Thailand. Some dental similarities with anthropoids were noted early on, such as deep jaws and wide basins that separate low molar cusps. The best known genera were Pondaungia and Amphipithecus (Ciochon and Gunnell 2002; see Figure 9.14). Another amphipithecid, Siamopithecus from Thailand, has very rounded molars and was probably a seed-eater (Figure 9.19). In addition to teeth and jaws, some cranial fragments, ankle material, and ends of postcranial bones have been found for Pondaungia. There are important resemblances between the postcranial bones of Pondaungia and those of adapoids, suggesting adapoid affinities for the amphipithecidae. This would imply that the resemblances with anthropoids in the teeth are convergent, based on similarities in diet (see Ciochon and Gunnell 2002). Unfortunately, the association between postcranial bones and teeth is not definite. With other primates in these faunas (including eosimiids), one cannot be certain that the postcranial bones belong with the teeth. Some researchers suggest that some bones belong to a sivaladapid (or asiadapid) and others to an early anthropoid (Beard et al. 2007; Marivaux et al. 2003). Additional well-associated material of amphipithecids would help to clear up this uncertainty.

Platyrrhine Dispersal to South America
Today there is an impressive diversity of primates in South and Central America. These are considered to be part of a single clade, the Platyrrhini. Primates colonized South America sometime in the Eocene from an African source. In the first half of the 20th century, the source of platyrrhines was a matter of major debate among paleontologists, with some favoring a North American origin (e.g., Simpson 1940).
Part of the reason for this debate is that South America was an island in the Eocene. Primates needed to cross open ocean to get there from either North America or Africa, although the distance from the former was shorter. Morphology yields clues to platyrrhine origins. The first known primates in South America have more in common morphologically with African primates than with North American ones. At the time, anthropoids were popping up in North Africa, whereas the only euprimates in North America were adapoids and omomyoids. Despite lacking a bony ear tube, early platyrrhines shared a great deal with other anthropoids, including full postorbital closure and fusion of the mandibular symphysis.
The means by which a population of small North African primates managed to disperse across the Atlantic and survive to colonize South America remains a mystery. The most plausible scenario is one of rafting. That is, primates must have been trapped on vegetation that was blown out to sea by a storm. The vegetation then became a sort of life raft, which eventually landed ashore, dumping its passengers in South America. Rodents probably arrived in South America in the same way (Antoine et al. 2012).
Once ashore, platyrrhines must have crossed South America fairly rapidly because the earliest-known primates from that continent are from Peru (Bond et al. 2015). Soon after that, platyrrhines were in Bolivia, namely Branisella. By the Miocene, platyrrhines were living in extreme southern Argentina and were exploiting a variety of feeding niches. The Early Miocene platyrrhines were all somewhat plesiomorphic in their morphology, but some features that likely arose by ecological convergence suggest (to some) relationships with extant platyrrhine families. This has led to a lively debate about the pattern of primate evolution in South America (Kay 2015; Kay and Fleagle 2010; Rosenberger 2010). By the Middle Miocene, clear representatives of modern families were present in a diverse fauna from La Venta, Colombia (Wheeler 2010). The Plio-Pleistocene saw the emergence of giant platyrrhines as well as several taxa of platyrrhines living on Caribbean islands (Cooke et al. 2016).
The story of platyrrhines seems to be one of amazing sweepstakes dispersal, followed by rapid diversification and widespread geographic colonization of much of South America. After that, dramatic extinctions resulted in the current, much-smaller geographic distribution of platyrrhines. These extinctions were probably caused by changing climates, leading to the contraction of forests. Platyrrhines dispersed to the Caribbean and to Central America, with subsequent extinctions in those regions that might have been related to interactions with humans. Unlike anthropoids of Africa and Asia, platyrrhines do not seem to have evolved any primarily terrestrial forms and so have always been highly dependent on forests.
Special Topic: Jonathan Perry and Primates of the Extreme South
Many primates are very vulnerable to ecological disturbance because they are heavily dependent on fruit to eat and trees to live in. This is one reason why so many primates are endangered today and why many of them went extinct due to climatic and vegetational changes in the past. I (Jonathan Perry) have conducted paleontological research focusing on primates that lived on the edge of their geographic distribution. This research has taken me to extreme environments in the Americas: southern Patagonia, the Canadian prairies, western Wyoming, and the badlands of eastern Oregon.
Santa Cruz Province in Argentina is as far south as primates have ever lived. The Santa Cruz fauna of the Miocene has yielded a moderate diversity of platyrrhines, each with slightly different dietary adaptations. These include Homunculus, first described by Florentino Ameghino in 1891 (Figure 9.20). Recent fieldwork by my colleagues and I in Argentina has revealed several skulls of Homunculus as well as many parts of the skeleton (Kay et al. 2012). The emerging profile of this extinct primate is one of a dedicated arboreal quadruped that fed on fruits and leaves. Many of the foods eaten by Homunculus must have been very tough and were probably covered and impregnated with grit; we suspect this because the cheek teeth are very worn down, even in young individuals, and because the molar tooth roots were very large, presumably to resist strong bite forces (Perry et al. 2010, 2014).

I began working in Argentina while a graduate student at Duke University. I participated as a field assistant in a team led by my Ph.D. advisor, Richard F. Kay, and Argentine colleagues Sergio F. Vizcaíno and M. Susana Bargo. Most of the localities examined belong to a suite of beach sites known since the 1800s and visited by many field parties from various museums in the early 1900s. Since 2003, our international team of paleontologists from the U.S. and Argentina has visited these localities every single year (Figure 9.21). Over time, new fossils and new students have led to new projects and new approaches, including the use of microcomputed tomography (microCT) to visualize and analyze internal structures of the skeleton.

Planet of Apes
Geologic Activity and Climate Change in the Miocene
The Miocene Epoch was a time of mammalian diversification and extinction, global climate change, and ecological turnover. In the Miocene, there was an initial warming trend across the globe with the expansion of subtropical forests, followed by widespread cooling and drying with the retreat of tropical forests and replacement with more open woodlands and eventually grasslands. It was also a time of major geologic activity. On one side of the globe, South America experienced the rise of the Andes Mountains. On the other side, the Indian subcontinent collided with mainland Asia, resulting in the rise of the Himalayan Mountains. In Africa, volcanic activity promoted the development of the East African Rift System. Critical to the story of ape evolution was the exposure of an intercontinental landbridge between East Africa and Eurasia, permitting a true planet of apes (Figure 9.22).

Geographic Distribution: Africa, Asia, Europe
The world of the Miocene had tremendous ape diversity compared to today. The earliest records of fossil apes are from Early Miocene deposits in Africa. However, something dramatic happened around 16 million years ago. With the closure of the ancient Tethys Sea, the subsequent exposure of the Gomphotherium Landbridge, and a period of global warming, the Middle–Late Miocene saw waves of emigration of mammals (including primates) out of Africa and into Eurasia, with evidence of later African re-entry for some (Harrison 2010). Some of the mammals that dispersed from Africa to Eurasia and back were apes. Though most of these early apes left no modern descendants, some of them gave rise to the ancestors of modern apes—including hominins (Figure 9.23).

Where Are the Monkeys? Diversity in the Miocene
Whereas the Oligocene deposits in the Fayum of Egypt have yielded the earliest-known catarrhine fossils, the Miocene demonstrates some diversification of Cercopithecoidea. However, compared to the numerous and diverse Miocene apes (see below), monkeys of the Miocene are very rare and restricted to a single extinct family, the Victoriapithecidae (Figure 9.24). This family contains the earliest definite cercopithecoids. These monkeys are found from northern and eastern Africa between 20 million and 12.5 million years ago (Miller et al. 2009). The best known early African monkey is Victoriapithecus (Figure 9.25), a small-bodied (approximately 7 kg; 15 lbs.), small-brained monkey. Bilophodonty, known to be a hallmark of molar teeth of modern cercopithecoid, was present to some extent in Victoriapithecids. Victoriapithecus has been reconstructed as being more frugivorous and perhaps spent more time on the ground (terrestrial locomotion) than in the trees (arboreal locomotion; Blue et al. 2006). The two major groups of cercopithecoids today are cercopithecines and colobines. The earliest records demonstrating clear members of each of these two groups are at the end of the Miocene. Examples include the early colobine Microcolobus from Kenya and the early cercopithecine Pliopapio from Ethiopia.
|
Family
|
Genera
|
Morphology
|
Location
|
Age1
|
|
Victoriapithecidae2 |
Victoriapithecus |
Long, sloping face. Round, narrowly spaced orbits. Deep cheek bones. Well-developed sagittal crest. |
Africa |
Early to Middle Miocene |
|
Proconsulidae3 |
Proconsul |
Short face. Generalized dentition. Arboreal quadruped. Probably tailless. |
Africa and Arabia |
Early to Middle Miocene |
|
Pongidae |
Gigantopithecus |
Largest primate ever. Deep jaws and low rounded molars. |
Asia |
Miocene to Present |
|
1 Derived from Fleagle 2013. 2 See Benefit and McCrossin 1997 and Fleagle 2013. 3 See Begun 2007. |
|
|||
The Story of Us, the Apes
African Ape Diversity
The Early Miocene of Africa has yielded around 14 genera of early apes (Begun 2003). Many of these taxa have been reconstructed as frugivorous arboreal quadrupeds (Kay 1977). One of the best studied of these genera is the East African Proconsul (Family Proconsulidae; see Figure 9.24). Several species have been described, with body mass reconstructions ranging from 17 to 50 kg (approximately 37–110 lbs.). A paleoenvironmental study reconstructed the habitat of Proconsul to be a dense, closed-canopy tropical forest (Michel et al. 2014). No caudal vertebrae (tail bones) have been found in direct association with Proconsul postcrania, and the morphology of the sacrum is consistent with Proconsul lacking a tail (Russo 2016; Ward et al. 1991).
Overall, the African ape fossil record in the Late Miocene is sparse, with seven fossil localities dating between eleven and five million years ago (Pickford et al. 2009). Nevertheless, most species of great apes live in Africa today. Where did the progenitors of modern African apes arise? Did they evolve in Africa or somewhere else? The paucity of apes in the Late Miocene of Africa stands in contrast to the situation in Eurasia. There, ape diversity was high. Furthermore, several Eurasian ape fossils show morphological affinities with modern hominoids (apes). Because of this, some paleoanthropologists suggest that the ancestors of modern African great apes recolonized Africa from Eurasia toward the end of the Miocene (Begun 2002). However, discoveries of Late Miocene hominoids like the Kenyan Nakalipithecus (9.9 million to 9.8 million years ago), the Ethiopian Chororapithecus (10.7 million to 10.1 million years ago), and the late-Middle Miocene Namibian Otavipithecus (13 million to 12 million years ago) fuel an alternative hypothesis—namely that African hominoid diversity was maintained throughout the Miocene and that one of these taxa might, in fact, be the last common ancestor of extant African apes (Kunimatsu et al. 2007; Mocke et al. 2002). The previously underappreciated diversity of Late Miocene apes in Africa might be due to poor sampling of the fossil record in Africa.
Eurasian Ape Diversity
With the establishment of the Gomphotherium Landbridge (a result of the closure of the Eastern Mediterranean seaway; Rögl 1999), the Middle Miocene was an exciting time for hominoid radiations outside of Africa (see Figure 9.23). Eurasian hominoid species exploited their environments in many different ways in the Miocene. Food exploitation ranged from soft-fruit feeding in some taxa to hard-object feeding in others, in part owing to seasonal fluctuations and the necessary adoptions of fallback foods (DeMiguel et al. 2014). For example, the molars of Oreopithecus bambolii (Family Hominidae) have relatively long lower-molar shearing crests, suggesting that this hominoid was very folivorous (Ungar and Kay 1995). Associated with variation in diet, there is great variation in the degree to which cranial features (e.g., zygomatic bone or supraorbital tori) are developed across the many taxa (Cameron 1997); however, Middle Miocene fossils tend to exhibit relatively thick molar enamel and relatively robust jaws (Andrews and Martin 1991).
In Spain, the cranium with upper dentition, part of a mandible, and partial skeleton of Pliobates (Family Pliobatidae), a small-bodied ape (4–5 kg; 9–11 lbs.), was discovered in deposits dating to 11.6 million years ago (Alba et al. 2015). It is believed to be a frugivore with a relative brain size that overlaps with modern cercopithecoids. The fossilized postcrania of Pliobates suggest that this ape might have had a unique style of locomotion, including the tendency to walk across the branches of trees with its palms facing downward and flexible wrists that permitted rotation of the forearm during climbing. However, the anatomy of the distal humerus differs from those of living apes in ways that suggest that Pliobates was less efficient at stabilizing its elbow while suspended (Benefit and McCrossin 2015). Two other recently described apes from Spain, Pierolapithecus and Anoiapithecus, are known from relatively complete skeletons. Pierolapithecus had a very projecting face and thick molar enamel as well as some skeletal features that suggest (albeit controversially) a less suspensory locomotor style than in extant apes (Moyà-Solà et al. 2004). In contrast to Pierolapithecus, the slightly younger Anoiapithecus has a very flat face (Moyà-Solà et al. 2009).
Postcranial evidence for suspensory or well-developed orthograde behaviors in apes does not appear until the Late Miocene of Europe. Primary evidence supporting these specialized locomotor modes includes the relatively short lumbar vertebrae of Oreopithecus (Figure 9.26) and Dryopithecus (Maclatchy 2004). Further, fossil material of the lower torso of O. bambolii (which dates to the Pan-hominin divergence) conveys a higher degree of flexion-extension abilities in the lumbar region (lower back) than what is possible in extant apes. Additionally, the hindlimb of O. bambolii is suggested to have supported powerful hip adduction during climbing (Hammond et al. 2020). The Late Miocene saw the extinction of most of the Eurasian hominoids in an event referred to as the Vallesian Crisis (Agustí et al. 2003). Among the latest surviving hominoid taxa in Eurasia were Oreopithecus and Gigantopithecus, the latter of which held out until the Pleistocene in Asia and was probably even sympatric with Homo erectus (Cachel 2015).

The Origins of Extant Apes
The fossil record of the extant apes is somewhat underwhelming: it ranges from being practically nonexistent for some taxa (e.g., chimpanzees) to being a little better for others (e.g., humans). There are many possible reasons for these differences in fossil abundance, and many are associated with the environmental conditions necessary for the fossilization of bones. One way to understand the evolution of extant apes that is not so dependent on the fossil record is via molecular evolutionary analyses. This can include counting up the differences in the genetic sequence between two closely related species to estimate the amount of time since these species shared a common ancestor. This is called a molecular clock, and it is often calibrated using fossils of known absolute age that stand in for the last common ancestor of a particular clade. Molecular clock estimates have placed the Hylobatidae and Hominidae split between 19.7 million and 24.1 million years ago, the African ape and Asian ape split between 15.7 million and 19.3 million years ago, and the split of Hylobatidae into its current genera between 6.4 million and 8 million years ago (Israfil et al. 2011).
Small Ape Origins and Fossils
Unfortunately, the fossil record for the small (formerly “lesser”) apes is meager, particularly in Miocene deposits. One possible early hylobatid is Laccopithecus robustus, a Late Miocene catarrhine from China (Harrison 2016). Although it does share some characteristics with modern gibbons and siamangs (including an overall small body size and a short face), Laccopithecus most likely represents a plesiomorphic stem catarrhine and is therefore distantly related to extant apes (Jablonski and Chaplin 2009). A more likely candidate for the hylobatid stem is another Late Miocene taxon from China, Yuanmoupithecus xiaoyuan. Interpretation of its phylogenetic standing, however, is complicated by contradicting dental features—some of them quite plesiomorphic—which some believe best place Yuanmoupithecus as a stem hylobatid (Harrison 2016). Recently, a Middle Miocene Indian fossil ape, Kapi ramnagarensis, has extended the fossil record of small apes by approximately five million years. Its teeth are suggestive of a shift to a more frugivorous diet and it is likely a stem hylobatid (Gilbert et al. 2020). The history of Hylobatidae becomes clearer in the Pleistocene, with fossils representing extant genera.
Great Ape Origins and Fossils
The most extensive fossil record of a modern great ape is that of our own genus, Homo. However, the evolutionary history of the Asian great ape, the orangutan (Pongo), is becoming clearer. Today, orangutans are found only on the islands of Borneo and Sumatra. However, Pleistocene-aged teeth, attributed to Pongo, have been found in Cambodia, China, Laos, Peninsular Malaysia, and Vietnam—demonstrating the vastness of the orangutan’s previous range (Ibrahim et al. 2013; Wang et al. 2014). Sivapithecus from the Miocene of India and Pakistan is represented by many specimens, including parts of the face. Sivapithecus is very similar to Pongo, especially in the face, and it probably is closely related to ancestral orangutans (Pilbeam 1982). Originally, jaws and teeth belonging to the former genus Ramapithecus were thought to be important in the origin of humans (Simons 1961), but now these are recognized as specimens of Sivapithecus (Kelley 2002). Postcranial bones of Sivapithecus, however, suggest a more generalized locomotor mode—including terrestrial locomotion—than seen in Pongo (Pilbeam et al. 1990). Stable carbon and oxygen isotope data from dental enamel have reconstructed the paleoecological space of Sivapithecus (as well as the contemporaneous Late Miocene pongine Khoratpithecus) within the canopies of forested habitats (Habinger et al. 2022).
A probable close relative of Sivapithecus is the amazing Gigantopithecus (see Figure 9.24). Known only from teeth and jaws from China and India (e.g., Figure 9.27), this ape probably weighed as much as 270 kg (595 lbs.) and was likely the largest primate ever (Bocherens et al. 2017). Because of unique features of its teeth (including molarized premolars and patterns of wear) and its massive size, it has been reconstructed as a bamboo specialist, somewhat like the modern panda. Small silica particles (phytoliths) from grasses have been found stuck to the molars of Gigantopithecus (Ciochon et al. 1990). Recent studies evaluating the carbon isotope composition of the enamel sampled from Gigantopithecus teeth suggest that this ape exploited a wide range of vegetation, including fruits, leaves, roots, and bamboo (Bocherens et al. 2017). Its face is reminiscent of that of modern orangutans and it might belong in the same family, Pongidae (Kelley 2002).

In Africa, the first fossil to be confidently attributed to Pan, and known to be the earliest evidence of a chimpanzee, was described based on teeth found in Middle Pleistocene deposits in the Eastern Rift Valley of Kenya (McBrearty and Jablonski 2005). Paleoenvironmental reconstructions of this locality suggest that this early chimpanzee was living in close proximity to early Homo in a closed-canopy wooded habitat. Similarly, fossil teeth and mandibular remains attributed to two species of Middle-Late Miocene apes—Chororapithecus abyssinicus (from Ethiopia; Suwa et al. 2007) and Nakalipithecus nakayamai (from Kenya; Kunimatsu et al. 2007)—have been suggested as basal members of the gorilla clade.
While the deposits of Eastern Africa have yielded a profound record of our fossil hominin ancestors, the continent’s rainforests remain a “palaeontological desert” (Rosas et al. 2022). Clearly, more work is needed to fill in the large gaps in the fossil record of the nonhuman great apes. The twentieth century witnessed the discovery of many hominin fossils in East Africa, which have been critical for improving our understanding of human evolution. While twenty-first-century conservationists fight to prevent the extinction of the living great apes, perhaps efforts by twenty-first-century paleoanthropologists will yield the evolutionary story of these, our closest relatives.
Summary
While there are large gaps in the fossil records linking primates to early hominins, evolutionary trends make it clear that humans are one branch of the broader primate family tree. In this chapter we go over the major development that characterize primate evolution: enhanced vision, grasping hands and feet, greater reliance on social behavior, and increased brain complexity. It is these traits which distinguish primates from other mammals and furthermore, help define major subsections within the primate class, such as strepsirrhines, haplorhines, monkeys, apes, and ultimately hominins.
Within this chapter, we also examine how anthropologists reconstruct these evolutionary relationships. Fossil evidence has provided key information about when and where different primates lived, while genetic data such as skeletal features allow researchers to understand how extinct species moved, ate, and interacted with their environments. Just as crucial are the influences of Earth’s changing environments: continental drift, glacial cycles, and long-term climate shifts have repeatedly reshaped habitats, driving both extinctions and the emergence of new adaptive forms, including the emergence of our own human lineage.
Review Questions
- Compare three major hypotheses about primate origins, making reference to each one’s key ecological reason for primate uniqueness.
- Explain how changes in temperature, rainfall, and vegetation led to major changes in primate biogeography over the Early Tertiary.
- List some euprimate features that plesiadapiforms have and some that they lack.
- Contrast adapoids and omomyoids in terms of life habits.
- Describe one piece of evidence for each of the adapoid, omomyoid, and tarsier origin hypotheses for anthropoids.
- Discuss the biogeography of the origins of African great apes and orangutans using examples from the Miocene ape fossil record.
Key Terms
Adapoidea: Order: Primates. One of the earliest groups of euprimates (true primates; earliest records from the early Eocene).
Adaptive radiations: Rapid diversifications of single lineages into many species which may present unique morphological features in response to different ecological settings.
Ancestral traits: Features that were inherited from a common ancestor and which remain (largely) unchanged.
Anthropoids:Group containing monkeys and apes, including humans.
Auditory bulla: The rounded bony floor of the middle ear cavity.
Bilophodonty: Dental condition in which the cusps of molar teeth form ridges (or lophs) separated from each other by valleys (seen, e.g., in modern catarrhine monkeys).
Catarrhines: Order: Primates; Suborder: Anthropoidea; Infraorder: Catarrhini. Group, with origins in Africa and Asia, that contains monkeys and apes, including humans.
Clade:Group containing all of the descendants of a single ancestor. A portion of a phylogenetic tree represented as a bifurcation (node) in a lineage and all of the branches leading forward in time from that bifurcation.
Convergent evolution: The independent evolution of a morphological feature in animals not closely related (e.g., wings in birds and bats).
Crown: Smallest monophyletic group (clade) containing a specified set of extant taxa and all descendants of their last common ancestor.
Diastema: Space between adjacent teeth.
Diffuse coevolution: The ecological interaction between whole groups of species (e.g., primates) with whole groups of other species (e.g., fruiting trees).
Ectotympanic: Bony ring or tube that holds the tympanic membrane (eardrum).
Euprimates: Order: Primates. True primates or primates of modern aspect.
Haplorhines: Group containing catarrhines, platyrrhines, and tarsiers.
Hominins: Modern humans and any extinct relatives more closely related to us than to chimpanzees.
Mandibular symphysis: Fibrocartilaginous joint between the left and right mandibular segments, located in the midline of the body.
Omomyoidea: Order: Primates; Superfamily: Omomyoidea. One of the earliest groups of euprimates (true primates; earliest record in the early Eocene).
Petrosal bone: The portion of the temporal bone that houses the inner ear apparatus.
Plagiaulacoid: Dental condition where at least one of the lower cheek-teeth (molars or premolars) is a laterally compressed blade.
Platyrrhines: Order: Primates; Suborder: Anthropoidea; Infraorder: Platyrrhini. Group containing monkeys found in the Americas.
Plesiadapiforms: Order: Plesiadapiformes. Archaic primates or primate-like placental mammals (Early Paleocene–Late Eocene).
Plesiomorphic: Having features that are shared by different groups which arose from a common ancestor.
Stem: Taxa that are basal to a given crown group but are more closely related to the crown group than to the closest living sister taxon of the crown group.
Strepsirrhines: Order: Primates; Suborder: Stresirrhini. Group containing lemurs, lorises, and galagos (does not include tarsiers).
Toothcomb: Dental condition found in modern strepsirrhines in which the lower incisors and canines are laterally compressed and protrude forward at a nearly horizontal inclination. This structure is used in grooming.
For Further Exploration
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Begun, David R. 2010. “Miocene Hominids and the Origins of the African Apes and Humans.” Annual Review of Anthropology 39: 67–84.
Fleagle, John G. 2013. Primate Adaptation and Evolution. Third edition. San Diego, CA: Academic Press.
Gebo, Daniel L., ed. 1993. Postcranial Adaptations in Nonhuman Primates. Dekalb: Northern Illinois University Press.
Godfrey, Laurie R., and William L. Jungers. 2002. “Quaternary Fossil Lemurs.” In The Primate Fossil Record, edited by Walter C. Hartwig, 97–121. Cambridge: Cambridge University Press.
Godinot, Marc. 2006. “Lemuriform Origins as Viewed from the Fossil Record.” Folia Primatologica 77 (6): 446–464.
Kay, Richard F. 2018. “100 Years of Primate Paleontology.” American Journal of Physical Anthropology 165 (4): 652–676.
Marivaux, Laurent. 2006. “The Eosimiid and Amphipithecid Primates (Anthropoidea) from the Oligocene of the Bugti Hills (Balochistan, Pakistan): New Insight into Early Higher Primate Evolution in South Asia.” Palaeovertebrata, Montpellier 34 (1–2): 29–109.
Martin, R. D. 1990. Primate Origins and Evolution: A Phylogenetic Reconstruction. Princeton: Princeton University Press.
Rose, Kenneth D., Marc Godinot, and Thomas M. Bown. 1994. “The Early Radiation of Euprimates and the Initial Diversification of Omomyidae.” In Anthropoid Origins: The Fossil Evidence, edited by John G. Fleagle and Richard F. Kay, 1–28. New York: Plenum Press.
Ross, Callum F. 1999. “How to Carry Out Functional Morphology.” Evolutionary Anthropology 7 (6): 217–222.
Seiffert, Erik R. 2012. “Early Primate Evolution in Afro-Arabia.” Evolutionary Anthropology: Issues, News, and Reviews 21(6): 239–253.
Szalay, Frederic S., and Eric Delson. 1979. Evolutionary History of the Primates. New York: Academic Press.
Ungar, Peter S. 2002. “Reconstructing the Diets of Fossil Primates.” In Reconstructing Behavior in the Primate Fossil Record, edited by Joseph Plavcan, Richard F. Kay, William Jungers, and Carel P. van Schaik, 261–296. New York: Kluwer Academic/Plenum Publishers.
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Acknowledgments
We are immensely grateful to the editors of this book, Drs. Beth Shook, Lara Braff, Katie Nelson, and Kelsie Aguilera, for their time and commitment to making this knowledge freely accessible to all, and for giving us the opportunity to participate in this important project.
Amanda Wolcott Paskey, M.A., Cosumnes River College
AnnMarie Beasley Cisneros, M.A., American River College
Student contributors to this chapter: Peyton Dagg, Bryana Henry, and Anoriel Jacques
This chapter is a revision from "Chapter 11: Archaic Homo” by Amanda Wolcott Paskey and AnnMarie Beasley Cisneros. In Explorations: An Open Invitation to Biological Anthropology, first edition, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under CC BY-NC 4.0.
Learning Objectives
- Identify the main groupings of Archaic Homo sapiens, such as Neanderthals.
- Explain how shifting environmental conditions required flexibility of adaptations, both anatomically and behaviourally.
- Describe the unique anatomical and cultural characteristics of Archaic Homo sapiens, including Neanderthals, in contrast to other hominins.
- Articulate how Middle Pleistocene hominin fossils fit into evolutionary trends including cranial capacity (brain size) development, cultural innovations, and migration patterns.
- Identify the shared traits, regional variations, and local adaptations among Archaic Homo sapiens.
- Detail the increased complexity and debates surrounding the classification of hominins in light of transitional species, species admixture, etc.
Breaking the Stigma of the "Caveman"
What do you think of when you hear the word “caveman”? Perhaps you imagine a character from a film such as The Croods, Tarzan, and Encino Man or from the cartoon The Flintstones. Maybe you picture the tennis-playing, therapy-going hairy Neanderthals from Geico Insurance commercials. Or perhaps you imagine characters from The Far Side or B.C. comics. Whichever you picture, the character in your mind is likely stooped over with a heavy brow, tangled long locks and other body hair, and clothed in animal skins, if anything. They might be holding a club with a confused look on their face, standing at the entrance to a cave or dragging an animal carcass to a fire for their next meal (see Figure 12.1). You might have even signed up to take this course because of what you knew—or expected to learn—about “cavemen.”

These images have long been the stigma and expectation about our ancestors at the transition to modern Homo sapiens. Tracing back to works as early as Carl Linnaeus, scientists once propagated and advanced this imagery, creating a clear picture in the minds of early scholars that informed the general public, even through today, that Archaic Homo sapiens, “cavemen,” were somehow fundamentally different and much less intelligent than we are now. Unfortunately, this view is overly simplistic, misleading, and incorrect. Understanding what Archaic Homo sapiens were actually like requires a much more complex and nuanced picture, one that comes into sharper focus as continuing research uncovers more about the lives of our not-too-distant (and not-too-different) ancestors.
The first characterizations of Archaic Homo sapiens were formed from limited fossil evidence in a time when ethnocentric and species-centric perspectives (anthropocentrism) were more widely accepted and entrenched in both society and science. Today, scientists are working from a more complete fossil record from three continents (Africa, Asia, and Europe), and genetic evidence informs their analyses and conclusions. The existence of Archaic Homo sapiens marks an exciting point in our lineage—a point at which many modern traits had emerged and key refinements were on the horizon. Anatomically, humans today are not that much different from Archaic Homo sapiens.
The Changing Environment
While modern climate change is of critical concern today due to its cause (human activity) and pace (unprecedentedly rapid), the existence of global climate change itself is not a recent phenomenon. The focus of this chapter, the Middle Pleistocene (roughly between 780 kya and 125 kya), is the time period in which Archaic Homo sapiens appears in the fossil record—a time that witnessed some of the most drastic climatic changes in human existence. During this time period, there were 15 major and 50 minor glacial events in Europe, alone.
What exactly is glaciation? When scientists talk about glacial events, they are referring to the climate being in an ice age. This means that the ocean levels were much lower than today, because much of the earth’s water was tied up in large glaciers or ice sheets. Additionally, the average temperature would have been much cooler, which would have better supported an Arctic or tundra-adapted plant-and-animal ecosystem in northern latitudes. The most interesting and relevant features of Middle Pleistocene glacial events are the sheer number of them and their repeated bouts: this era alternated between glacial periods and warmer periods, known as interglacials. In other words, the planet wasn’t in an ice age the whole time.
You can see the dramatic and increasing fluctuations in temperature, recorded through foraminifera, in Figure 12.2. The distance between highs and lows demonstrates the severity of temperature shifts. Much as the Richter scale represents more intense earthquakes with more dramatic peaks, so too does this chart, which uses dramatic peaks to demonstrate intense temperature swings.

Glacial periods are defined by Earth’s average temperature being lower. Worldwide, temperatures are reduced, with cold areas becoming even colder. Huge portions of the landscape may have become inaccessible during glacial events due to the formation of glaciers and massive ice sheets. In Europe, the Scandinavian continental glacier covered what is today Ireland, England, Sweden, Norway, Denmark, and some of continental Europe. Plant and animal communities shifted to lower latitudes along the periphery of ice sheets. Additionally, some new land was opened during glacials. Evaporation with little runoff reduced sea levels by as much as almost 150 metres, shifting coastlines outward by in some instances as much as almost 100 kilometres. Additionally, land became exposed that connected what were previously unconnected continents such as Africa into Yemen at the Gulf of Aden.
Glacial periods also affected equatorial regions and other regions that are today thought of as warmer or at least more temperate parts of the globe, including Africa. While these areas were not covered with glaciers, increased global glaciation resulted in lower sea levels and expanded coastlines. Cooler temperatures were accompanied by the drying of the climate, which caused significantly reduced rainfall, increased aridity, and the expansion of deserts. It is an interesting question to consider whether the same plants and animals that lived in these regions prior to the ice ages would be able to survive and thrive in this new climate. Plant and animal communities shifted in response to the changing climate, whenever possible.
Surviving During the Middle Pleistocene
Rather than a single selective force, the Middle Pleistocene was marked by periods of fluctuation, not just cold periods. Interglacials interrupted glaciations, reversing trends in sea level, coastline, temperature, precipitation, and aridity, as well as glacier size and location. Interglacials are marked by increased rainfall and a higher temperature, which causes built-up ice in glaciers to melt. This leads to glacial retreat, which is the shrinking of glaciers and the movement of the glaciers back toward the poles, as we’ve seen in our lifetime. During interglacials, sea levels increase, flooding some previously exposed coastlines and continental connections. In addition, plant and animal communities shift accordingly, often finding more temperate climates to the north and less arid and more humid climates in the tropics (Van Andel and Tzedakis 1996).
Scientists have found that the Olorgesailie region in southern Kenya was at various times in the Middle Pleistocene a deep lake, a drought-dried lakebed with an area criss-crossed by small streams, and a grassland. While various animal species would have moved in and out of the area as the climate shifted, some animal species went extinct, and new, often related, species took up residence. The trend, scientists noted, was that animals with more specialized features went extinct and animals with more generalized features, such as animals we see today, survived in this changing climatic time period. For example, a zebra with specialized teeth for eating grass was ultimately replaced by a zebra that could eat both grass and other types of vegetation. If this small, localized example shows such a dramatic change in terms of the environment and the plant and animal biocommunities, what would have been the impact on humans?
There is no way humans could have escaped the effects of Middle Pleistocene climate change, no matter what region of the world they were living in. As noted earlier, and as evidenced by what was seen in the other biotic communities, humans would have faced changing food sources as previous sources of food may have gone extinct or moved to a different latitude. Depending on where they were living, fresh water may have been limited. Durial glacials, lower sea levels would have given humans more land to live on, while the interglacials would have reduced the available land through the increase in rainfall and associated sea level rise. Dry land connections between the continents would have made movement from one continent to another by foot easier at times than today, although these passageways were not consistently available through the Middle Pleistocene due to the glacial/interglacial cycle. Finally, as evidenced by the Olorgesailie region in Kenya, during the Middle Pleistocene animal species that were overly specialized to one particular type of environment were less likely to survive when compared to their more generalized counterparts. Evidence suggests that this same pattern may have held true for Archaic Homo sapiens, in terms of their ability to survive this dramatic period of climate change.
Defining Characteristics of Archaic Homo sapiens
Archaic Homo sapiens share our species name but are distinguished by the term “Archaic” as a way of recognizing both the long period of time between their appearance and ours, as well as the way in which human traits have continued to evolve over time—making Archaic Homo sapiens look slightly different from us today, despite being considered the same species. Living throughout Africa, and the Middle East during the Middle Pleistocene, Archaic Homo sapiens are considered, in many ways, transitional between Homo erectus and modern Homo sapiens (see Figure 12.3). Archaic Homo sapiens share the defining trait of an increased brain size of at least 1,100 cc and averaging 1,200 cc, although there are significant regional and temporal (time) variations. Because of these variations, scientists disagree on whether these fossils represent a single, variable species or multiple, closely related species (sometimes called Homo antecessor, Homo bodoensis, Homo heidelbergensis, Homo georgicus, Homo neanderthalensis, and Homo rhodesiensis).
An active area of scholarship in the discipline involves reconciling the diversity of species from this time period and establishing the phylogenetic relationships between them. The term “Archaic Homo sapiens” can mean different things to different scholars within the discipline. The intent of this chapter is to provide an understanding of the diversity of this time period and provide data used to make interpretations from among the most likely possibilities. Although we recognize that some anthropologists split many of these fossils into separate species, until the issue is resolved at the discipline level, this chapter will rely on the widely used naming conventions that refer to many fossils from this time period as Archaic Homo sapiens. We will discuss these contemporaneous fossils as a unit, with the exception of a particularly unique population living in Europe and West Asia known as the Neanderthals, which we will examine separately.
|
Trait |
Homo erectus |
Archaic Homo sapiens (including Neaderthals) |
Anatomically Modern Homo sapiens |
|
Time |
1.8 mya–200,000 ya |
600,000–40,000 ya |
315,000 ya–today |
|
Brain size |
900 cc |
1,200 cc (1,500 cc when including Neanderthals) |
1,400 cc |
|
Skull Shape |
Long and low, angular |
Intermediate |
Short and high, globular |
|
Forehead |
Absent |
Emerging |
Present |
|
Nasal Region |
Projecting nasal bones (bridge of the nose), no midfacial prognathism |
Wider nasal aperture and some midfacial prognathism, particularly pronounced among Neanderthals |
Narrower nasal aperture, no midfacial prognathism |
|
Chin |
Absent |
Absent |
Present |
|
Other Facial Features |
Large brow ridge and large projecting face |
Intermediate |
Small brow ridge and retracted face |
|
Other Skull Features |
Nuchal torus, sagittal keel, thick cranial bone |
Projecting occipital bone, often called occipital bun in Neanderthals; intermediate thickness of cranial bone |
Small bump on rear of skull, if anything; thin cranial bone |
|
Dentition |
Large teeth, especially front teeth |
Slightly smaller teeth; front teeth still large; retromolar gap in Neanderthals |
Smaller teeth |
|
Postcranial Features |
Robust bones of skeleton |
Robust bones of skeleton |
More gracile bones of skeleton |
When comparing Homo erectus, Archaic Homo sapiens, and anatomically modern Homo sapiens, one can see that Archaic Homo sapiens are intermediate in their physical form. For some features, this follows the trends first seen in Homo erectus with other features having early, less developed forms of traits seen in modern Homo sapiens. For example, Archaic Homo sapiens trended toward less angular and higher skulls than Homo erectus. However, the archaic skulls were not as short and globular and had less developed foreheads compared to anatomically modern Homo sapiens. Archaic Homo sapiens had smaller brow ridges and a less-projecting face than Homo erectus and slightly smaller teeth, although incisors and canines were often about as large as those of Homo erectus. Archaic Homo sapiens also had a wider nasal aperture, or opening for the nose, and a forward-projecting midfacial region, which is later seen more fully developed among Neanderthals and is known as midfacial prognathism. The occipital bone often projected and the cranial bone was of intermediate thickness, somewhat reduced from Homo erectus but not nearly as thin as that of anatomically modern Homo sapiens. The postcrania remained fairly robust. Identifying a set of features that is unique to Archaic Homo sapiens is a challenging task, due to both individual and geographic variation—these developments were not all present to the same degree in all individuals. Neanderthals are the exception, as they had several unique traits that made them notably different from modern Homo sapiens as well as their closely related Archaic cousins.

The one thing that is clear about Archaic Homo sapiens is that, despite general features, there is a lot of regional variation, which is first seen in the different Homo erectus specimens across Asia and Africa. While the general features of Archaic Homo sapiens, identified earlier, are present in the fossils of this time period, there are significant regional differences. The majority of this regional variation lies in the degree to which fossils have features more closely aligned with Homo erectus or with anatomically modern Homo sapiens.

To illustrate this point, we will examine three exemplary specimens, one from each of the three continents on which Archaic Homo sapiens lived. First, in Africa, a specimen from Broken Hill is one of several individuals found in the Kabwe lead mine in Zambia. It had a large brain (1,300 cc) and taller cranium as well as many Homo erectus-like skull features, including massive brow ridges, a large face, and thick cranial bones (Figure 12.4). Second, one partial crania from Dali, China, is representative of Archaic Homo sapiens in Asia, with large and robust features with heavy brow ridges, akin to what is seen in Homo erectus, and a large cranial capacity intermediate between Homo erectus and anatomically modern Homo sapiens (Figure 12.5). Third, an almost-complete skeleton was found in northern Spain at Atapuerca. Atapuerca 5 (Figure 12.6) has thick cranial bone, an enlarged cranial capacity, intermediate cranial height, and a more rounded cranium than seen previously. Additionally, Atapuerca 5 demonstrates features that foreshadow Neanderthals, including increased midfacial prognathism. After examining some of the fossils such as those from Kabwe, Dali, and Atapuerca, the transitional nature of Archaic Homo sapiens is clear: their features place them squarely between Homo erectus and modern Homo sapiens.

Due to the transitional nature of Archaic Homo sapiens, identifying the time period with which they are associated is problematic and complex. Generally, it is agreed that Archaic Homo sapiens lived between 600,000 and 200,000 years ago, with regional variation and overlap between Homo erectus on the early end of the spectrum and modern Homo sapiens and Neanderthals on the latter end. The earliest-known Archaic Homo sapiens fossils tentatively date to about 600,000 years ago in Africa, to around 300,000 years ago in Asia, and to about 350,000 years ago in Europe (and potentially as early as 600,000 years ago). Determining the end point of Archaic Homo sapiens is also problematic since it largely depends upon when the next subspecies of Homo sapiens appears and the classification of highly intermediate specimens. For example, in Africa, the end of Archaic Homo sapiens is met with the appearance of modern Homo sapiens, while in Europe it is the appearance of Neanderthals that is traditionally seen as marking the transition from other Archaic Homo sapiens.
It is important to remember that this time period is represented by many branching relationships and assuming an evolutionary trajectory that follows a single linear path would not be correct. Even still, Archaic Homo sapiens mark an important chapter in the human lineage, connecting more ancestral forms, such as Homo erectus, to modern Homo sapiens. During this period of climatic transition and fluctuation, Archaic Homo sapiens mirror the challenges of their environments. Showing increasing regional variation due to the need for local adaptation, there is no single archetype for this group; the defining characteristic seems to be variability.
Neanderthals
One well-known population of Archaic Homo sapiens are the Neanderthals, named after the site where they were first discovered in the Neander Valley, or “thal” in German, located near Dusseldorf, Germany. Popularly known as the stereotypical “cavemen” examined at the outset of this chapter, recent research is upending long-held beliefs about this group of Archaics. Neanderthal behaviour was increasingly complex, far beyond what was exhibited by even other Archaic Homo sapiens discussed throughout this chapter. We implore you to forget the image of the iconic caveman and have an open mind when exploring the fossil evidence of the Neanderthals.
It is important to understand why Neanderthals are separated from other Archaic Homo sapiens. Unlike the rest of Archaic Homo sapiens, Neanderthals are easily defined and identified in many ways. Evidence suggests the time period when Neanderthals lived was between 150,000 and 40,000 years ago. There is a clear geographic boundary of where Neanderthals lived: western Europe, the Middle East, and western Asia. No Neanderthal fossils have ever been discovered outside of this area, including Africa. This is a bit curious, as other Archaics seem to have adapted in Africa and then migrated elsewhere, but Neanderthals’ regional association makes sense in light of the environment to which they were best adapted: namely, extreme cold weather. Additionally, Neanderthals have a unique and distinct cluster of physical characteristics. While a few aspects of Neanderthals are shared among some Archaic Homo sapiens, such as the types of tools, most Neanderthal anatomical and behavioural attributes are unique to them.
Neanderthals lived during some of the coldest times during the last Ice Age and at far northern latitudes. This means Neanderthals were living very close to the glacial edge, rather than in a more temperate region of the globe like some of their Archaic Homo sapiens relatives. While able to survive in arctic conditions, most Neanderthal sites dating to the glacial periods were found farther away from the severe cold, in a steppe tundra-like environment, which would have been more hospitable to Neanderthals, and their food sources, both flora and fauna (Ashton 2002; Nicholson 2017; Richter 2006). Their range likely expanded and contracted along with European glacial events, moving into the Middle East during glacial events when Europe became even cooler, and when the animals they hunted would have moved for the same reason. During interglacials, when Europe warmed a bit, Neanderthals and their prey would have been able to move back into Western Europe. Clearly, the true hallmark of Neanderthals is their adaptation to an unstable environment, shifting between warm and cold, as the climate was in constant flux throughout their existence (Adler Et al. 2003; Boettger Et al. 2009).
Many of the Neanderthals’ defining physical features are more extreme and robust versions of traits seen in other Archaic Homo sapiens, clustered in this single population. Brain size, namely an enlargement of the cranial capacity, is one such trait. The average Neanderthal brain size is around 1,500 cc, and the range for Neanderthal brains can extend to upwards of 1,700 cc. The majority of the increase in the brain occurs in the occipital region, or the back part of the brain, resulting in a skull that has a large cranial capacity with a distinctly long and low shape that is slightly wider than previous forms at the far back of the skull. Modern humans have a brain size comparable to that of Neanderthals; however, our brain expansion occurred in the frontal region of the brain, not the back, as in Neanderthal brains. This difference is also the main reason why Neanderthals lack the vertical forehead that modern humans possess. They simply did not need an enlarged forehead, because their brain expansion occurred in the rear of their brain. Due to cranial expansion, the back of the Neanderthal skull is less angular (as compared to Homo erectus), but not as rounded as Homo sapiens, producing an elongated shape, akin to a football.
Another feature that continues the trend noted in previous hominins is the enlargement of the nasal region, or the nose. Neanderthal noses are large and have a wide nasal aperture, which is the opening for the nose. While the nose is only made up of two bones, the nasals, the true size of the nose can be determined by looking at other facial features, including the nasal aperture, and the angle of the nasal and maxillary, or facial bones. In Neanderthals, these indicate a large, forward-projecting nose that appears to be pulled forward away from the rest of the face. This feature is further emphasized by the backward-sloping nature of the cheekbones, or the zygomatic arches. The unique shape and size of the Neanderthal nose is often characterized by the term midfacial prognathism—a jutting out of the middle portion of the face, or nose. This is in sharp contrast to the prognathism exhibited by other hominins, who exhibited prognathism, or the jutting out, of their jaws.
The teeth of the Neanderthals follow a similar pattern seen in the Archaic Homo sapiens, which is an overall reduction in size, especially as compared to the extremely large teeth seen in the genus Australopithecus. However, while the teeth continued to reduce, the jaw size did not keep pace, leaving Neanderthals with an oversized jaw for their teeth, and a gap between their final molar and the end of their jaw. This gap is called a retromolar gap.
The projecting occipital bone present in other Archaic Homo sapiens is also more prominent in Neanderthals, extending the trend found in Archaics. Among Neanderthals, this projection of bone is easily identified by its bun shape on the back of the skull and is known as an occipital bun. This projection appears quite similar to a dinner roll in size and shape. Its purpose, if any, remains unknown.
Continuing the Archaic Homo sapiens trend, Neanderthal brow ridges are prominent but somewhat smaller in size than those of Homo erectus and earlier Archaic Homo sapiens. In Neanderthals, the brow ridges are also often slightly less arched than those of other Archaic Homo sapiens.
In addition to extending traits present in Archaic Homo sapiens, Neanderthals possess several distinct traits. Neanderthal infraorbital foramina, the holes in the maxillae or cheek bones through which blood vessels pass, are notably enlarged compared to other hominins. The Neanderthal postcrania are also unique in that they demonstrate increased robusticity in terms of the thickness of bones and body proportions that show a barrel-shaped chest and short, stocky limbs, as well as increased musculature. These body portions are seen across the spectrum of Neanderthals—in men, women, and children.
Traditionally, many of the unique traits that Neanderthals possess were seen as adaptations to the extreme cold, dry environments in which they often lived and which exerted strong selective forces. For example, Bergmann’s and Allen’s Rules dictate that an increased body mass and short, stocky limbs are common in animals that live in cold conditions. Neanderthals were said to have matched the predictions of Bergmann’s and Allen’s Rules perfectly (Churchill 2006). In addition, the Neanderthal skull also exhibits adaptations to the cold. Neanderthals’ large infraorbital foramina allow for larger blood vessels, increasing the volume of blood that is found closest to the skin, which helps to keep the skin warmer. Their enlarged noses resulted in longer nasal passages and mucus membranes that warmed and moistened cold air before it reached the lungs. The Neanderthals’ larger nose has long been thought to have acted as a humidifier, easing physical exertion in their climate, although research on this particular trait continues to be studied and debated (Rae Et al. 2011).
New research, however, seems to suggest that these unique skeletal adaptations might not have been strict adaptations to cold weather (Evteev Et al. 2017; Pearce Et al. 2013). For example, large brow ridges might have served as a way to shade the face from the sun. The increased occipital portion of the brain, some researchers state, was to support a larger visual system present in Neanderthals. This visual system would have given them increased light sensitivity, which would have been useful in higher latitudes that had dark winters. And, while recent modeling of nostril airflow on reconstructed Neanderthal specimens supports the notion that Neanderthals had extensive mucus membranes inside their noses, the data shows that modern Homo sapiens are superior to Neanderthals in our ability to use our noses as a way to warm and cool air. However, Neanderthals were able to snort air through their noses better than we can. Why is this important? When combined with the fact that Neanderthals tended to prefer a more temperate, tundra-like environment, and that other physical traits suggest that Neanderthals had huge bodies that needed massive amounts of calories to sustain them, the picture gets clearer. Massive amounts of energy would have been required to power a Neanderthal body, and anything that might have made them more calorically efficient would have been favoured. Efficient breathing, through larger noses into large lungs, meaning deeper breaths, would have been favoured. To further save energy expenditure, body sizes might have been sacrificed as well. These same types of adaptations are similar to ones seen in children today who are born in high altitudes, not cold climates. Figure 12.7 provides a summary of these unique features of the Neanderthal.
|
Distinct Neanderthal Anatomical Features |
|
|
Brain Size |
1,500 cc average |
|
Skull Shape |
Long and low |
|
Brow Ridge Size |
Large |
|
Nose Size |
Large, with midfacial prognathism |
|
Dentition |
Reduced, but large jaw size, creating retromolar gap |
|
Occipital Region |
Enlarged occipital region, occipital bun |
|
Other Unique Cranial Features |
Large infraorbital foramina |
|
Postcranial Features |
Short and stocky body, increased musculature, barrel-shaped chest |
A classic example of a Neanderthal with all of the characteristics mentioned above is the nearly complete La Ferrassie 1 Neanderthal, from France. This is a male skeleton, with a brain size of around 1640cc, an extremely large nose and infraorbital foramina, brow ridges that are marked in size, and an overall robust skeleton (Figure 12.8).

Neanderthal Culture: Tool Making and Use
One key Neanderthal adaptation was their cultural innovations, which are an important way that hominins adapt to their environment. As you recall, Homo erectus's tools, Acheulean handaxes, represented an increase in complexity over Oldowan tools, allowing more efficient removal of meat and possibly calculated scavenging. In contrast, Neanderthal tools mark a significant innovation in tool-making technique and use with Mousterian tools (named after the Le Moustier site in southwest France). These tools were significantly smaller, thinner, and lighter than Acheulean handaxes and formed a true toolkit. The materials used for Mousterian tools were of higher quality, which allowed for both more precise toolmaking and tool reworking when the tools broke or dulled after frequent reuse. The use of higher-quality materials is also indicative of required forethought and planning to acquire them for tool manufacture. It has been suggested that the Neanderthals, unlike Homo erectus, saved and reused their tools, rather than making new ones each time a tool was needed.

Mousterian tools are constructed in a very unique manner, utilizing the Levallois technique (Figure 12.9), named after the first finds of tools made with this technique, which were discovered in the Levallois-Perret suburb of Paris, France. The Levallois technique is a multistep process that requires preparing the core, or raw material, in a specific way that will yield flakes that are roughly uniform in dimension. The flakes are then turned into individual tools. The preparation of the core is akin to peeling a potato or carrot with a vegetable peeler—when peeling vegetables, you want to remove the skin in long, regular strokes, so that you are taking off the same amount of the vegetable all the way around. In the same way, the Levallois technique requires removing all edges of the cortex, or outside surface of the raw material, in a circle before removing the lid. The flakes, which will eventually be turned into the individual tools, can then be removed from the core. The potential yield of tools from one core would be many, as seen in Figure 12.10, compared to all previous tool-making processes, in which one core yielded a single tool. This manufacturing process might be considered the ultimate zero-waste tool-making technique (Delpiano Et al. 2018).

It is suggested that Neanderthal tools were used for a variety of purposes, including cutting, butchering, woodworking or antler working, and hide working. Additionally, because the Mousterian tools were lighter than previous stone tools, Neanderthals could haft, or attach the tool onto a handle, as the stone would not have been too heavy (Degano Et al. 2019). Neanderthals attached small stone blades onto short wood or antler handles to make knives or other small weapons, as well as attached larger blades onto longer shafts to make spears. New research examining tar-covered stones and black lumps at several Neanderthal sites in Europe suggests that Neanderthals may have been making tar by distilling it from birch tree bark, which could have been used to glue the stone tool onto its handle. If Neanderthals were, in fact, manufacturing tar to act as glue, this would predate modern humans in Africa using tree resin or similar adhesives by nearly 100,000 years.
Evidence shows that raw materials used by Neanderthals came from distances as far away as 100 km. This could indicate a variety of things regarding Neanderthal behaviour, including a limited trade network with other Neanderthal groups or simply a large area scoured by Neanderthals when collecting raw materials. While research on specific applications continues, it should be clear from this brief discussion that Neanderthal tool manufacturing was much more complex than previous tool-making efforts, requiring technical expertise, patience, and skills beyond toolmaking to carry out.
Neanderthal Culture: Hunting and Diet
With their more sophisticated suite of tools and robust muscular bodies, Neanderthals were better armed for hunting than previous hominins. The animal remains in Neanderthal sites show that, unlike earlier Archaic Homo sapiens, Neanderthals were very effective hunters who were able to kill their own prey, rather than relying on scavenging. Though more refined than the tools of earlier hominins, the Neanderthal spear was not the kind of weapon that would have been thrown; rather, it would have been used in a jabbing fashion (Churchill 1998; Kortlandt 2002). This may have required Neanderthals to hunt in groups rather than individually and made it necessary to approach their prey quite closely (Gaudzinski-Windheuser Et al. 2018). Remember, the animals living with Neanderthals were very large-bodied due to their adaptations to cold weather; this would have included species of deer, horses, and bovids (relatives of the cow).
Isotopes from Neanderthal bones show that meat was a significant component of their diet, similar to that seen in carnivores like wolves (Bocherens Et al. 1999; Jaouen Et al. 2019; Richards Et al. 2000). In addition to large prey, their diet included ibex, seals, rabbits, and pigeons. Though red meat was a critical component of the Neanderthal diet, evidence shows that at times they also ate limpets, mussels, and pine nuts. Tartar examined from Neanderthal teeth in Iraq and Belgium reveal that they also ate plant material including wheat, barley, date palms, and tubers, first cooking them to make them palatable (Henry Et al. 2010). While Neanderthals’ diet varied according to the specific environment in which they lived, meat comprised up to 80% of their diet (Wiẞin Et al. 2015).
Neanderthal Culture: Caring for the Injured and Sick
While the close-range style of hunting used by Neanderthals was effective, it also had some major consequences. Many Neanderthal skeletons have been found with significant injuries, which could have caused paralysis or severely limited their mobility. Many of the injuries are to the head, neck, or upper body. Thomas Berger and Erik Trinkaus (1995) conducted a statistical comparative analysis of Neanderthal injuries compared to those recorded in modern-day workers’ compensation reports and found that the closest match was between Neanderthal injuries and those of rodeo workers. Rodeo professionals have a high rate of head and neck injuries that are similar to the Neanderthals’ injuries. What do Neanderthals and rodeo workers have in common? They were both getting very close to large, strong animals, and at times their encounters went awry.
The extensive injuries sustained by Neanderthals are evident in many fossil remains. Shanidar 1 (Figure 12.11), an adult male found at the Shanidar site in northern Iraq and dating to 45,000 ya, has a lifetime of injuries recorded in his bones (Stewart 1977). Shanidar 1 sustained—and healed from—an injury to the face that would have likely caused blindness. His lower right arm was missing and the right humerus shows severe atrophy, likely due to disuse. This pattern has been interpreted to indicate a substantial injury that required or otherwise resulted in amputation or wasting away of the lower arm. Additionally, Shanidar 1 suffered from bony growths in the inner ear that would have significantly impaired his hearing and severe arthritis in the feet. He also exhibited extensive anterior tooth wear, matching the pattern of wear found among modern populations who use their teeth as a tool. Rather than an anomaly, the type of injuries evident in Shanidar 1 are similar to those found in many other Neanderthal fossils, revealing injuries likely sustained from hunting large mammals as well as demonstrating a long life of physical activity.


The pattern of injuries is as significant as the fact that Shanidar 1 and other injured Neanderthals show evidence of having survived their severe injuries. One of the earliest-known Neanderthal discoveries—the one on whom misinformed analysis shaped the stereotype of the species for nearly a century—is the La Chapelle-aux-Saints Neanderthal (Trinkaus 1985). The La Chapelle Neanderthal had a damaged eye orbit that likely caused blindness and suffered arthritis of the spine (Dawson and Trinkaus 1997). He had also lost most of his teeth, many of which he had lived without for so long that the mandibular and maxillary bones were partially reabsorbed due to lack of use. The La Chapelle Neanderthal was also thought to be at least in his mid-forties at death, an old age for the rough life of the Late Pleistocene—giving rise to his nickname, “the Old Man.” To have survived so long with so many injuries that obviously precluded successful large game hunting, he must have been taken care of by others. Such caretaking behaviour is also evident in the survival of other seriously injured Neanderthals, such as Shanidar 1. Long thought to be a hallmark modern human characteristic, taking care of the injured and elderly, for example preparing or pre-chewing food for those without teeth, indicates strong social ties among Neanderthals.
Neanderthal Culture: Ritual Life
Such care practices may have been expressed upon death as well. Nearly complete Neanderthal skeletons are not uncommon in the fossil record, and most are well preserved within apparently deliberate burials that involve deep graves and bodies found in specific, often fetal or flexed positions (Harrold 1980). Discoveries of pollen in a grave at the Shanidar site in the 1960s led scientists to think that perhaps Neanderthals had placed flowering plants in the grave, an indication of ritual ceremony or spirituality so common in modern humans. But more recent investigations have raised some doubt about this conclusion (Pomeroy Et al. 2020). The pollen may have been brought in by burrowing rodents. Claims of grave goods or other ornamentation in burials are similarly debated, although possible.
Some tantalizing evidence for symbolism, and debatably, ritual, is the frequent occurrence of natural pigments, such as ochre (red) and manganese dioxide (black) in Neanderthal sites that could have been used for art. However, the actual uses of pigments are unclear, as there is very little evidence of art or paintings in Mousterian sites. One exception may be the recent discovery in Spain of a perforated shell that appears to be painted with an orange pigment, which may be evidence of Neanderthal art and jewelry. However, many pigments also have properties that make them good emulsifiers in adhesive (like for attaching a stone tool to a wooden handle) or useful in tanning hides. So the presence of pigment may or may not be associated with symbolic thought; however, it definitely does show a technological sophistication beyond that of earlier Archaic hominins.
Dig Deeper: Evidence of Endocannibalism Among the Neanderthals
Krapina, a Neanderthal site in Croatia, has recently sparked new archeological discourse as many investigations upon fossilized remains show evidence of post-mortem modifications and manipulations of limbs and bones through the use of tools (Rougier Et al., 2016). These findings provide compelling evidence of Neanderthals engaging in cannibalism as part of their post-mortem practices. Additionally, these uncovered remains were found to have died of natural causes as opposed to being killed, showing that even the earliest humans may have had some sort of morality or ethics surrounding the cannibalizing of their kin, meaning they were aware of death in a social context as opposed to merely a physical one.
While the original evidence in Krapina was uncovered in 1901, Croatian geologist and paleontologist Dragutin Gorjanović-Kramberger’s discovery of fragmented and burned human bones (Ullrich, 2005) was not yet confirmed to be linked to endocannibalism until much later. Whereas the discovery of burned bones does not mean they were being prepared for consummation, due to its context among other findings, this information supports the hypothesis that early hominids conducted post-mortem rituals and practices with their dead. Building on Gorjanović-Kramberger’s research, Herbert Ullrich wrote in Anthropologie (2005) that broken bones—resulting from post-mortem bodily manipulations—were “defleshed in preparation for secondary burial” (2005, 251) and intentionally left outside rock shelters, while selectively chosen bones were seemingly brought inside for use in mortuary practices.
For nearly 150 years, since the first Neanderthal skeletal remains were discovered, anthropologists and researchers have continued to debate the cognitive, social, and physical abilities of this species. In 2016, Rougier and colleagues wrote in Scientific Reports, furthering the research, presenting 99 new Neanderthal remains found in Goyet, Belgium. Among these remains, similar evidence of human-induced alterations was identified, including signs of butchering, consumption, and the use of bones to modify stone tools (Rougier Et al., 2016). This discovery provides significant support for the presence of cannibalistic behaviour among Northern European Neanderthals, contributing to the growing body of evidence that Neanderthals engaged with death in ways that reflect social awareness, ritual behaviour, and complex cultural practices.
Contemporary Cases of Prion Disease Related to Endocannibalism
The evidence of endocannibalism does not end with early hominids; with Australian medical anthropologists recording thousands of cannibalism-related prion disease occurrences present in populations up until 2009 (Radford & Scragg, 2013). Following a mysterious epidemic of a new form of spongiform encephalopathy–a “progressive degenerative disease of the central nervous system” (2013, p.29)–anthropological research regarding the the cultural mortuary rites within the Okapa region of Papua New Guinea have linked the newfound disease ‘Kuru’ to post-mortem consumption of human remains (Radford & Scragg, 2013; Collinge Et al., 2006).
Local oral histories collected during the first investigations by these researchers in the 1950s traced the earliest cases back to the 1920s, with detailed case histories. Epidemiological data revealed a strong correlation between the spread of Kuru and participation in mortuary feasts, in which the deceased were ritually consumed as part of funerary rites (2006, p. 2070). From 1957 to 2004, over 2,700 cases were reported, with mortality peaking at over 200 deaths annually in the late 1950s (2006, p.2070); however, following the cessation of cannibalism in the easly 1960s due to governmental efforts, natural transmission of the disease has stopped, dropping the death toll dramatically, with the “last three single cases reported in 2005, 2007, and 2009” (Radford & Scragg, p.48).
The Lasting Gift of Neanderthals: Tantalizing New Directions for Research
Examining the more recent time period in which Neanderthals lived and the extensive excavations completed across Europe allows for a much more complete archaeological record from this time period. Additionally, the increased cultural complexity such as complex tools and ritual behaviours expressed by Neanderthals left a more detailed record than previous hominins. Intentional burials enhanced preservation of the dead and potentially associated ritual behaviours. Such evidence allows for a more complete and nuanced picture of this species.

Additional analyses are possible on many Neanderthal finds, due to increased preservation of bone, the amount of specimens that have been uncovered, and the recency in which Neanderthals lived. We should be cautious, however, to consider the potential bias of many Neanderthal sites. Overwhelmingly, Neanderthal skeletons are found complete, with injuries or evidence of disease in caves. Does this mean all Neanderthals lived a tough, disease-wrought life? Probably not. It does, however, indicate that the sick were cared for by others, and that they lived in environments that preserved their bodies incredibly well. These additional studies include the examination of dental calculus and even DNA analysis. While limited, samples of Neanderthal DNA have been successfully extracted and analysed. Research thus far has identified specific genetic markers that show some Neanderthals were light-skinned and probably red-haired with light eyes. Genetic analyses, different from the typical hominin reconstruction done with earlier species, allow scientists to further investigate soft tissue markers of Neanderthals and other more recent hominin species. These studies offer striking conclusions regarding Neanderthal traits, their physical appearance, and their culture, as reflected in these artists’ reconstructions (Figure 12.12).

Dr. Svante Pääbo (Figure 12.13), of the Max Planck Institute for Evolutionary Anthropology, has been at the forefront of much of this new research, largely in the form of genomic studies (The Nobel Prize 2022). Awarded the Nobel Prize for Physiology or Medicine in 2022, Pääbo is known primarily for his work with ancient DNA. He has successfully sequenced mitochondrial DNA (mtDNA) as well as the entire Neanderthal genome from nuclear DNA. His genomic work has led to the realization that Denisovans are genetically distinct from Neanderthals, as well as the recent identification of a Neanderthal father and teenage daughter, which he discovered by looking for unique DNA markers in the fossil record. Additionally, Pääbo’s genomic work has provided researchers with additional lines of evidence regarding the connections between hominin fossils (such as Neanderthals) and modern people, their time of divergence, and current genetic overlap. The work of Pääbo has even formalized a new field of study within anthropology—paleogenomics. To stay up to date with Dr. Svante Pääbo’s work, be sure to follow his lab’s website.
Neanderthal Culture: Communicating through Speech
To successfully live in groups and to foster cultural innovations, Neanderthals would have required at least a basic form of communication in order to function, possibly using a speech-based communication system. The challenge with this line of research is that speech, of course, is not preserved, so indirect evidence must be used to support this conclusion. It is thought that Neanderthals would have possessed some basic speech, as evidenced from a variety of sources, including throat anatomy and genetic evidence (Lieberman 1971). There is only one bone in the human body that could demonstrate if a hominin was able to speak, or produce clear vocalizations like modern humans, and that is the hyoid, a U-shaped bone that is found in the throat and is associated with the ability to precisely control the vocal cords. Very few hyoid bones have been found in the archaeological record; however, a few have been uncovered in Neanderthal burials. The shape of the Neanderthal hyoid is nearly identical to that of modern humans, pointing to the likelihood that they had the same vocal capabilities as modern humans. In addition, geneticists have uncovered a mutation present in both modern humans and Neanderthals—the FOXP2 gene—that is possibly linked to the ability to speak. However, other scientists argue that we cannot make sweeping conclusions that the FOXP2 gene accounts for speech due to small sample size. Finally, scientists have also pointed to the increasingly complex cultural behaviour of Neanderthals as a sign that symbolic communication, likely through speech, would have been the only way to pass down the skills needed to make, for example, a Levallois blade or to position a body for intentional burial.
Neanderthal Intelligence
One of the enduring questions about Neanderthals centres on their intelligence, specifically in comparison to modern humans. Brain volume indicates that Neanderthals certainly had a large brain, but it continues to be debated if Neanderthals were of equal intelligence to modern humans. Remember, creatures with larger body sizes tend to have larger brains; however, scaling of the brain is not always associated with greater intelligence (Alex 2018). Brain volume (cranial capacity), cultural complexity, tool use, and compassion toward their kind all point to an increase in intellect among Neanderthals when compared to previous hominins.
Yet, new research is suggesting additional differences between Neanderthal brains and our own. For example, Euluned Pearce and colleagues (2013), from the University of Oxford, noted the frontal lobes of Neanderthals and modern humans are almost identical. However, Neanderthals had a larger visual cortex—the portion of the brain involved in processing visual information. This would have left Neanderthals with less brain tissue for other functions, including those that would have aided them in dealing with large social groupings, one of the differences that has been suggested to exist between Neanderthals and modern humans. Other differences were found when geneticist John Blangero, from the Texas Biomedical Research Institute, compared data from the Neanderthal genome against data from modern study participants. Blangero and his colleagues (Blangero Et al. 2014) discovered that some Neanderthal brain components were very different, and smaller, than those in the modern sample. Differences were found in areas associated with the processing of information and controlling emotion and motivation, as well as overall brain connectivity. In short, as Blangero stated, “Neanderthals were certainly cognitively adept,” although their specific abilities may have differed from modern humans’ in key areas (qtd. in Wong 2015). This point has been echoed in other recent genetic studies comparing Neanderthal and anatomically modern human brains (el-Showk 2019).
Finally, scientists are fairly certain that Neanderthal brain development after birth was not the same as that of modern humans. After birth, anatomically modern Homo sapiens babies go through a critical period of brain expansion and cognitive development. It appears that Neanderthal babies’ brains and bodies did not follow the same developmental pattern (Smith Et al. 2010; Zollikofer and Ponce de León 2013). Modern humans enjoy an extended period of childhood, which allows children to engage in imaginative play and develop creativity that fosters cognitive skills. Neanderthals had a more limited childhood, with less development of the creative mind that may have affected their species’ success (Nowell 2016).
The exact nature of Neanderthal intelligence remains under investigation, however. Some studies disagree with the idea that Neanderthal intelligence had limitations compared to our own, noting the extensive evidence of Neanderthals having limb asymmetry. Their tools also have wear marks indicating that they were hand-dominant. This is further supported by marks on Neanderthal teeth that demonstrate hand dominance. The Neanderthal “stuff-and-cut method” of eating, noted by David Frayer and colleagues (Frayer Et al. 2012), would have seen Neanderthals hold a piece of meat in their teeth, while pulling it taut with one hand, and then using the other hand, their dominant one, to cut the meat off of the larger slab being held in their teeth. When looking at 17 Neanderthals and their tooth wear, only two do not show markings associated with a right-hand dominant individual eating in this manner. Further, it has been established that favouring the right hand is a key marker between modern humans and chimpanzees, and that handedness also relates to language development, in the form of bilateral brain development. That Neanderthals likely were hand-dominant suggests they had an indicator of bilateral brain development and a precondition for human speech.
The Middle Stone Age: Neanderthal Contemporaries in Africa
While Neanderthals made their home on and adapted to the European and Asian continents, evidence of fossil humans in Africa show they were also adapting to their local environments. These populations in Africa exhibit many more similarities to modern humans than Neanderthals, as well as overall evolutionary success. While the African fossil sample size is smaller and more fragmentary than the number of Neanderthal specimens across Europe and Asia, the African sample is interesting in that it represents a longer time period and larger geographical area. This group of fossils—often represented by the name “Middle Stone Age,” or MSA—dates to between 300,000 and 30,000 years ago across the entire continent of Africa. As with Archaic Homo sapiens, there is much variability seen in this African set of fossils. There are also a few key consistent elements: none of them exhibit Neanderthal skeletal features; instead, they demonstrate features that are increasingly consistent with anatomically modern Homo sapiens.
Similarities to Neanderthals and MSA contemporaries in Africa are seen, however, in their behavioural adaptations, including stone tools and other cultural elements. The tools associated with the specimens living in Africa during this time period are, like their physical features, varied. In some parts of Africa, namely Northern Africa, stone tools from this time so closely resemble Neanderthal tools that they are classified as Mousterian. In sub-Saharan Africa, the stone tools associated with these specimens are labeled as MSA. Some scholars argue that these could also be a type of Mousterian tools, but they are still typically subdivided based on geographical location.
Recall that Mousterian tools were much more advanced than their Acheulean predecessors in terms of how the stone tools were manufactured, the quality of the stones used, and the ultimate use of the tools that were made. In addition, recent evidence suggests that MSA tools may also have been heat treated—to improve the quality of the stone tool produced (Stolarczyk and Schmidt 2018). Evidence for heat treating is seen not only through advanced analysis of the tool itself but also through the residue of fires from this time period. Fire residues show a shift over time from small, short fires fuelled by grasses (probably intended for cooking) to larger, more intensive fires that required the exploitation of dry wood, exactly the type of fire that would have been needed for heat treating stone tools (Esteban Et al. 2018).
Other cultural elements seen with MSA specimens include the use of marine (sea-based) resources for their diet (Parkington 2003), manufacture of bone tools, use of adhesive and compound tools (e.g., hafted tools), shell bead production, engraving, use of pigments (such as ochre), and other more advanced tool-making technology (e.g., microlithics). While many of these cultural elements are also seen to a limited extent among Neanderthals, developments at MSA sites appear more complex. This MSA cultural expansion may have been a response to climate change or an increased use of language, complex communication, and/or symbolic thought. Others have suggested that the MSA cultural expansion was due to the increase of marine resources in their diet, which included more fatty acids that may have aided their cognitive development. Still others have suggested that the increased cultural complexity was due to increased interaction among groups, which spurred competition to innovate. Recent studies suggest that perhaps the best explanation for the marked cultural complexity of MSA cultural artifacts is best explained by the simple fact that they lived in diverse habitats (Kandel Et al. 2015). This would have necessitated a unique set of cultural adaptations for each habitat type (for example, specialized marine tools would have been needed along coastal sites but not at inland locations). Simply put, the most useful adaptation of MSA was their flexibility of behaviour and adaptability to their local environment. As noted previously in this chapter, flexibility of behaviour and physical traits, rather than specialization, seems to be a feature that was favoured in hominin evolution at this time.
Where Did They Go? The End of Neanderthals
While MSA specimens were increasingly successful and ultimately transitioned into modern Homo sapiens, Neanderthals disappeared from the fossil record by around 40,000 years ago. What happened to them? We know, based on genetics, that modern humans come largely from the modern people who occupied Africa around 300,000 to 100,000 years ago, at the same time that Neanderthals were living in northern Europe and Asia. As you will learn in Chapter 13, modern humans expanded out of Africa around 150,000 years ago, rapidly entering areas of Europe and Asia inhabited by Neanderthals and other Archaic hominins. Despite intense interest and speculation in fictional works about possible interactions between these two groups, there is very little direct evidence of either peaceful coexistence or aggressive encounters. It is clear, though, that these two closely related hominins shared Europe for thousands of years, and recent DNA evidence suggests that they occasionally interbred (Fu Et al. 2015). Geneticists have found traces of Neanderthal DNA (as much as 1% to 4%) in modern humans of European and Asian descent not present in modern humans from sub-Saharan Africa. This is indicative of limited regional interbreeding with Neanderthals.
While some interbreeding likely occurred, as a whole, Neanderthals did not survive. What is the cause for their extinction? This question has fascinated many researchers and several theories have been suggested, including:
- At the time that Neanderthals were disappearing from the fossil record, the climate went through both cooling and warming periods—each of which posed challenges for Neanderthal survival (Defleur and Desclaux 2019; Staubwasser Et al. 2018). It has been argued that as temperatures warmed, large-bodied animals, well adapted to cold weather, moved farther north to find colder environments or faced extinction. A shifting resource base could have been problematic for continued Neanderthal existence, especially as additional humans, in the form of modern Homo sapiens, began to appear in Europe and compete for a smaller pool of available resources.
- It has been suggested that the eruption of a European volcano 40,000 years ago could have put a strain on available plant resources (Golovanova Et al. 2010). The eruption would have greatly affected local microclimates, reducing the overall temperature enough to alter the growing season.
- Possible differences in cognitive development may have limited Neanderthals in terms of their creative problem solving. As much as they were biologically specialized for their environment, the nature of their intelligence might not have offered them the creative problem-solving skills to innovate ways to adapt their culture when faced with a changing environment (Pearce Et al. 2013).
- CRISPR gene-editing technology has been used in studies to evaluate potential differences between human and Neanderthal brains, based on differences in the genetic code. Potential differences include a Neanderthal propensity for mutations related to brain development that could account for more rapid brain development, maturation, synapse misfires, and less-orderly neural processes (Mora-Bermúdez Et al. 2022; Trujillo Et al. 2021). Fundamental differences in brain function at the cellular level may account for the differential survival rates of Neanderthal and modern human populations.
- There is evidence that suggests reproduction may have posed challenges for Neanderthals. Childbirth was thought to have been at least as difficult for female Neanderthals as anatomically modern Homo sapiens (Weaver and Hublin 2009). Female Neanderthals may have become sexually mature at an older age, even older than modern humans. This delayed maturation could have kept the Neanderthal population size small. A recent study has further suggested that male Neanderthals might have had a genetic marker on the Y chromosome that could have caused incompatibility between the fetus and mother during gestation; this would have had severe consequences for birth rate and survival (Mendez Et al. 2016). Even a small but continuous decrease in fertility would have been enough to result in the extinction of Neanderthals (Degioanni Et al. 2019).
- As mentioned above, the end of Neanderthal existence overlaps with modern human expansion into northern Europe and Asia. There is no conclusive direct evidence to indicate that Neanderthals and modern humans lived peacefully side by side, nor that they engaged in warfare, but by studying modern societies and the tendencies of modern humans, it has been suggested that modern humans may not have warmly embraced their close but slightly odd-looking cousins when they first encountered them (Churchill Et al. 2009). Nevertheless, direct competition with modern humans for the same resources may have contributed to the Neanderthals’ decline (Gilpin Et al. 2016); it may also have exposed them to new diseases, brought by modern humans (Houldcroft and Underdown 2016), which further decimated their population. Estimates of energy expenditures suggest Neanderthals had slightly higher caloric needs than modern humans (Venner 2018). When competing for similar resources, the slightly greater efficiency of modern humans might have helped them experience greater success in the face of competition—at a cost to Neanderthals.
It has been suggested that the Neanderthal populations were fairly small to begin with (estimated between 5,000 and 70,000 individuals; Bocquet-Appel and Degioanni 2013), one or a combination of these factors could have easily led to their demise. As more research is conducted, we will likely get a better picture of exactly what led to Neanderthal extinction.
Denisovans

While Neanderthals represent one regionally adapted branch of the Archaic Homo sapiens family tree, recent discoveries in Siberia and the Tibetan Plateau surprised paleoanthropologists by revealing yet another population that was contemporary with Archaic Homo sapiens, Neanderthals, and modern Homo sapiens. The genetic analysis of a child’s finger bone (Figure 12.14) and an adult upper third molar (Figure 12.15) from Denisova Cave in the Altai Mountains in Siberia by a team including Svante Pääbo discovered that the mitochondrial and nuclear DNA sequences reflected distinct genetic differences from all known Archaic populations. Dubbed “Denisovans” after the cave in which the bones were found, this population is more closely related to Neanderthals than modern humans, suggesting the two groups shared an ancestor who split from modern humans first, then the Neanderthal-Denisovan line diverged more recently (Reich Et al. 2010).

Denisovans share up to 5% of their DNA with modern Melanesians, aboriginal Australians, and Polynesians, and 0.2% of their DNA with other modern Asian populations and Native Americans. Additional studies have suggested one (Vernot Et al. 2018) or two (Browning Et al. 2018) separate points of time when interbreeding occurred between modern humans and Denisovans.
Genetic analysis reveals that Denisovans (potentially three distinct populations) had adaptations for life at high altitudes that prevented them from developing altitude sickness and hypoxia in extreme environments such as Tibet, where the average annual temperature is close to 0℃ and the altitude is more than a kilometre (about 4,000 feet) above sea level. Through protein analysis of a jawbone, one study (Chen Et al. 2019) has placed Denisovans in Tibet as early as 160,000 years ago. Genetic evidence of interbreeding has linked modern Tibetan populations with Denisovans 30,000 to 40,000 years ago, which implies that the unique high-altitude adaptations seen in modern Tibetans may have originated with Denisovans (Huerta-Sanchez Et al. 2014).
Other research suggests tantalizing new directions regarding Denisovans. Stone tools similar to those found in Siberia have been uncovered in the Tibetan plateau suggesting a connection between the Denisovan populations in those two areas (Zhang Et al. 2018). The molar of a young girl, possibly Denisovan, has been found in Laos and shows strong similarities to specimens from China (Demeter Et al. 2022). And DNA sequencing from discoveries in the Denisova Cave have yielded a genome that has been interpreted as the first-generation offspring of a Denisovan father and Neanderthal mother (Slon Et al. 2018). While this research is not yet conclusive and is still being interpreted, exciting new possibilities are being revealed. To stay up-to-date with new discoveries, consider following organizations such as the Smithsonian’s Human Origins Program on social media.
Pleistocene Hominins in East Asia: Homo longi and Yunxian
New fossil discoveries, along with advanced molecular analysis, are prompting re-evaluations of the complex evolutionary dynamics of non-sapiens Homo in the middle-late Pleistocene.
Homo longi
The Harbin cranium, often referred to as Homo longi, represents one of the most remarkable Middle Pleistocene human fossils discovered in East Asia. It is currently carbon-dated at 146 ka, allowing recent scholarship to use the Harbin cranium to argue for a distinct clade of hominins that evolved independently in Eastern Asia, separate from Neanderthals and modern humans (Ni et al., 2021). This clade is argued to include Dali, Jinniushan, Hualongdong, Xiahe, and Harbin, based on phylogenetic analyses using parsimony criteria and Bayesian tip-dating, forming a third East Asian lineage and a sister group to the Homo sapiens lineage.
The Harbin cranium exhibits a unique combination of traits, likely representing a species of Homo distinct from other Middle-to-Late Pleistocene taxa, including H. sapiens, H. Neanderthalensis, and H. Heidelbergensis/H. Rhodesiensis. Its overall size is exceptional, setting it apart from nearly all other known fossils; however, the proportions of the cranial vault align closely with those of other large late archaic Homo species, with an estimated brain volume of approximately 1,420 mL. Despite its considerable breadth, the face is relatively low in height and exhibits a zygomaxillary shape similar to that seen in H. sapiens and H. antecessor, a feature also observed in other Middle Pleistocene Chinese fossils from Dali and Jinniushan. Additionally, the face is attached to the braincase with reduced prognathism, forming a more perpendicular angle as in modern humans, in contrast to the pronounced midfacial prognathism of Neanderthals.
Yunxian, China: Possible cladistic origins
In 2025, researchers released findings that expanded the diversity and relationships of Homo lineages in the Pleistocene, based on an analysis of fossils found in 1989/90. Two skull fossils were found near the Hanjiang River in Hubei province, China, and were named the Yunxian 1 and Yunxian 2 crania—A third cranium has recently been discovered in the same sediment layer as the two previous finds (Tu et al., 2026). The first 2 specimens were in poor condition, fragmented and distorted; however, using recently developed techniques such as high-resolution computed tomography (CT) scans, image segmentation, and digital repositioning of fragments, the researchers incorporated elements from the Yunxian 1 crania with the less damaged Yunxian 2 cranium. The Yunxian 2 cranium is a mosaic of primitive and derived traits. It retains ancient characteristics seen in earlier hominins, a thick supraorbital torus, a broad palate, a long and low vault in lateral view, and a receding frontal contour; its frontal lobe remains low and narrow, similar to H. erectus fossils. Conversely, it lacks the strongly angulated occipital bone found in H. erectus as well as the protruding occipital region or central suprainiac fossa typical of Neanderthals. The big surprise is that the zygomaxillary region is transversely flat and faces anteriorly, a feature comparable to the Hualongdong and Harbin fossils and even H. sapiens. This intermediate morphology places Yunxian 2 in a distinct position, linking archaics like H. erectus with later Asian hominins such as Harbin, Dali, and Jinniushan. Phylogenetic and morphometric analyses confirm that Yunxian 2 represents an early member of the Asian H.longi Clade. The reconstructed Yunxian 2 cranium represents a major achievement in paleoanthropology, digitally restoring a badly damaged fossil from beyond the Middle Pleistocene. In fact, recent discussion suggests an even earlier date for the Yunxian fossils at approximately 1.77 Ma (Tu et al., 2026). In their analysis, they identify the Yunxian fossils as H.erectus, and the site as Early Pleistocene. Over 500 artifacts have been recovered from the site (cores, choppers, and flakes) typical of the Early Paleolithic. Nevertheless, the mosaic morphology of the Yunxian skulls reveals characteristics within the range of both H. erectus and the Harbin skull (among others), which may position these finds as being very close to the origins and diversification of the longi clade, as well as preserving transitional features close to the sapiens clade (Feng et al., 2025, p.1323).
The expanding fossil and genomic record of Archaic Homo has profoundly reshaped our understanding of Pleistocene human evolution. Discoveries, from an increasing variety of fossils found in eastern Asia, to advances in genomic research, demonstrate that multiple contemporaneous lineages once coexisted, interacted, diverged and, at times, interbred. These findings challenge earlier linear models of human ancestry and instead point to a dynamic evolutionary landscape marked by regional diversification, intermittent contact, and complex patterns of gene flow.
How Do These Fit In? Homo naledi and Homo floresiensis
Recently, some fossils have been unearthed that have challenged our understanding of the hominin lineage. The fossils of Homo naledi and Homo floresiensis are significant for several reasons but are mostly known for how they don’t fit the previously held patterns of hominin evolution. While we examine information about these species, we ask you to consider the evidence presented in this chapter and others to draw your own conclusions regarding the significance and placement of these two unusual fossil species in the hominin lineage.
Homo naledi

In 2013 recreational spelunkers uncovered a collection of bones deep in a cave network in Johannesburg, South Africa. The cave system, known as Rising Star, had been well documented by other cavers; however, it appears few people had ever gone as far into the cave as these spelunkers did. Lee Berger, paleoanthropologist at University of Witwatersrand, in Johannesburg, immediately put out a call for what he termed “underground astronauts” to begin recovery and excavation of the fossil materials. Unlike other excavations, Berger and most other paleoanthropologists would not be able to access the elusive site, as it was incredibly difficult to reach, and at some points there was only eight inches of space through which to navigate. The underground astronauts, all petite women anthropologists, were the only ones who were able to access this remarkable site. Armed with small excavation tools and a video camera, which streamed the footage up to the rest of the team at the surface, the team worked together and uncovered a total of 1,550 bones, representing at least 15 individuals, as seen in Figure 12.16. Later, an additional 131 bones, including an almost-complete cranium, were found in a nearby chamber of the cave, representing three more individuals (Figure 12.17). Berger called in a team of specialists to participate in what was dubbed “Paleoanthropology Summer Camp.” Each researcher specialized in a different portion of the hominin skeleton. With various specialists working simultaneously, more rapid analysis was possible of Homo naledi than most fossil discoveries.

While access to the site, approximately 80 m from any known cave entrance or opening, was treacherous for researchers, it must have been difficult for Homo naledi as well. The route included moving through a portion that is just 25 cm wide at some points, known as “Superman’s Crawl.” The only way to get through this section is by crawling on your stomach with one arm by your side and the other raised above your head. Past Superman’s Crawl, a jagged wall known as the Dragon’s Back would have been very difficult to traverse. Below that, a narrow vertical chute would have eventually led down to the area where the fossils were discovered. While geology changes over time and the cave system likely has undergone its fair share, it is not likely that these features arose after Homo naledi lived (Dirks Et al. 2017). This has made scientists curious as to how the bones ended up in the bottom of the cave system in the first place. It has been suggested that Homo naledi deposited the bones there, one way or another. If Homo naledi did deposit the bones, either through random disposal or intentional burial, this raises questions regarding their symbolic behaviour and other cultural traits, including the use of fire, to access a very dark cave system. Another competing idea is that a few individuals may have entered the cave system to escape a predator and then got stuck. To account for the sheer number of fossils, this would have had to happen multiple times.
The features of Homo naledi are well-documented due to the fairly large sample, which represents individuals of all sexes and a wide range of ages. The skull shape and features are very much like other members of the genus Homo—including a sagittal keel and large brow, like Homo erectus, and a well-developed frontal lobe, similar to modern humans—yet the brain size is significantly smaller than its counterparts, at approximately 500 cc (560 cc for males and 465 cc for females). The teeth also exhibit features of later members of the genus Homo, such as Neanderthals, including a reduction in overall tooth size. Homo naledi also had unique shoulder anatomy and curved fingers, indicating similarities to tree-dwelling primates, which is very different from any other hominin yet found. Perhaps the greatest shock of all is that Homo naledi has been dated to 335,000 to 236,000 years ago, placing it as a contemporary to modern Homo sapiens, despite its very primitive features. An additional specimen of a child, found in 2021, not only shares many of the unique features found in the adult specimen but will also add insight into the growth and development of individuals of this species (Brophy Et al. 2021).
Homo floresiensis
In a small cave called Liang Bua, on the island of Flores, in Indonesia, a small collection of fossils were discovered beginning in 2003 (Figure 12.18). The fossil fragments represent as many as nine individuals, including a nearly complete female skeleton. The features of the skull are very similar to that of Homo erectus, including the presence of a sagittal keel, an arching brow ridges and nuchal torus, and the lack of a chin (Figure 12.19). Homo floresiensis, as the new species is called, had a brain size that was remarkably small at 400 cc, and recent genetic studies suggest a common ancestor with modern humans that predates Homo erectus.


The complete female skeleton, who was an adult, was approximately a metre tall and would have weighed just under 30 kg, which is significantly shorter and just a few kilograms more than the average, modern, young elementary-aged child. A reconstructed comparison between an anatomically modern human and Homo floresiensis can be seen in Figure 12.20. The small size of the fossil has earned the species the nickname “the Hobbit.” Many questions have been asked about the stature of this species, as all of the specimens found also show evidence of diminutive stature and small brain size. Some explanations include pathology; however, this seems unlikely as all fossils found thus far demonstrate the same pattern. Another possible explanation lies in a biological phenomena seen in other animal species also found on the island, which date to a similar time period. This phenomenon, called insular dwarfing, is due to limited food resources on an island, which can create a selective pressure for large-bodied species to be selected for smaller size, as an island would not have been able to support their larger-bodied cousins for a long period of time. This phenomenon is the cause of other unique species known to have lived on the island at the same time, including the miniature stegodon, a dwarf elephant species.

There is ongoing research and debate regarding Homo floresiensis’ dates of existence, with some researchers concluding that they lived on Flores until perhaps as recently as 17,000 years ago, although they are more often dated to 100,000 to 60,000 years ago. Stone tools from that time period uncovered at the site are similar to other hominin stone tools found on the island of Flores. Homo floresiensis would have hunted a wide range of animals, including the miniature stegodon, giant rats, and other large rodents. Other animals on the island that could have threatened them include the giant komodo dragon. An interesting note about this island chain is that ancestors of Homo floresiensis would have had to traverse the open ocean in order to get there, as the nearest island is almost 10 km away, and there is little evidence to support that a land bridge connecting mainland Asia or Australia to the island would have been present. This separation from the mainland would also have limited the number of other animals, including predators and human species, that would have had access to the island. Anatomically modern Homo sapiens arrived on the island around 30,000 years ago and, if some researchers’ later dates for Homo floresiensis are correct, both species may have lived on Flores at the same time. The modern population living on the island of Flores today believes that their ancestors came from the Liang Bua cave; however, recent genetic studies have determined they are not related to Homo floresiensis (Tucci Et al. 2018).
Hominin Species Summaries
|
Hominin |
Archaic Homo sapiens |
|
Dates |
600,000–200,000 years ago (although some regional variation) |
|
Region(s) |
Africa, Europe, and Asia |
|
Famous discoveries |
Broken Hill (Zambia), Atapuerca (Spain) |
|
Brain size |
1,200 cc average |
|
Dentition |
Slightly smaller teeth in back of mouth, larger front teeth |
|
Cranial features |
Emerging forehead, no chin, projecting occipital region |
|
Postcranial features |
Robust skeleton |
|
Culture |
Varied regionally, but some continue to use Acheulean handaxe, others adopt Mousterian tool culture |
|
Other |
Lots of regional variation in this species |
|
Species |
Homo naledi |
|
Dates |
335,000–236,000 years ago |
|
Region(s) |
South Africa |
|
Famous discoveries |
Rising Star Cave |
|
Brain size |
500 cc average |
|
Dentition |
Reduced tooth size |
|
Cranial features |
Sagittal keel, large brow, well-developed frontal region |
|
Postcranial features |
Suspensory shoulder |
|
Culture |
unknown |
|
Other |
N/A |
|
Hominin |
Neanderthals |
|
Dates |
150,000–40,000 years ago |
|
Region(s) |
Western Europe, Middle East, and Western Asia only |
|
Famous discoveries |
Shanidar (Iraq), La Chapelle-aux-Saints (France) |
|
Brain size |
1500 cc average |
|
Dentition |
Retromolar gap |
|
Cranial features |
Large brow ridge, midfacial prognathism, large infraorbital foramina, occipital bun |
|
Postcranial features |
Robust skeleton with short and stocky body, increased musculature, barrel chest |
|
Culture |
Mousterian tools often constructed using the Levallois technique |
|
Other |
N/A |
|
Species |
Homo floresiensis |
|
Dates |
100,000–60,000 years ago, perhaps as recently as 17,000 years ago |
|
Region(s) |
Liang Bua, island of Flores, Indonesia |
|
Famous discoveries |
“The Hobbit” |
|
Brain size |
400 cc average |
|
Dentition |
unknown |
|
Cranial features |
Sagittal keel, arching brow ridges, nuchal torus, no chin |
|
Postcranial features |
Very short stature (approximately 3.5 ft.) |
|
Culture |
Tools similar to other tools found on the island of Flores |
|
Other |
N/A |
|
Hominin |
Denisovans |
|
Dates |
100,000–30,000 years ago |
|
Region(s) |
Siberia |
|
Famous discoveries |
Child’s finger bone and adult molar |
|
Brain size |
unknown |
|
Dentition |
Large molars (from limited evidence) |
|
Cranial features |
unknown |
|
Postcranial features |
unknown |
|
Culture |
unknown |
|
Other |
Closely related to Neanderthals (genetically) |
Summary
Research presented in the chapter contributes to why scientists have taken to nicknaming this time period “the muddle in the middle.” We know that the Middle Pleistocene picks up from Homo erectus and ends with the appearance of anatomically modern Homo sapiens. While the start and the end are clear, it’s the middle that is messy. As more research is conducted and more data is collected, rather than clarifying our understanding of the hominin lineage during this time period, it only inspires more questions, particularly about the relationships between hominins during this time period, including the oft-misunderstood Neanderthal. Research is painting a more detailed picture of Neanderthal intelligence and both biological and behavioural adaptations. At the same time, their relationship to other Middle Pleistocene hominins, including Denisovans, as well as modern humans, remains unclear.
Homo naledi and Homo floresiensis are clear outliers when compared to their contemporary hominin species. Each has surprised paleoanthropologists for both their archaic traits in relatively modern times and their unique combination of traits seen in archaic species and modern humans. While these finds have been exciting, they have also completely upended the assumed trajectory of the human lineage, causing scientists to re-examine assumptions about hominin evolution and what it means to be modern. Add this to the developments being made using ancient DNA, other new fossil discoveries, and other innovations in paleoanthropology, and you see that our understanding of Archaic Homo sapiens and others living during this time period is rapidly developing and changing. This is a true testament to the nature of science and the scientific method.
Clearly, hominins of the Middle Pleistocene are distinct from our species today. Yet, understanding the hominins that directly preceded our species and clarifying the evolutionary relationships between us is important to better understanding our own place in nature.
Review Questions
- What physical and cultural features are unique to Archaic Homo sapiens? How are Archaic Homo sapiens different in both physical and cultural characteristics from Homo erectus?
- Describe the specific changes to the brain and skull first seen in Archaic Homo sapiens. Why does the shape of the skull change so dramatically from Homo erectus?
- What role did the shifting environment play in the adaptation of Archaic Homo sapiens, including Neanderthals? Discuss at least one physical feature and one cultural feature that would have assisted these groups in surviving the changing environment.
- What does the regional variation in Archaic Homo sapiens represent in terms of the broader story of our species’ evolution?
- Describe the issues raised by the discoveries of Homo naledi and Homo floresiensis in the understanding of the story of the evolution of Homo sapiens.
Key Terms
Allele: Each of two or more alternative forms of a gene that arise by mutation and are found at the same place on a chromosome.
Anthropocentrism: A way of thinking that assumes humans are the most important species and leads to interpreting the world always through a human lens. Species-centric science and thought.
Cortex: The outside, or rough outer covering, of a rock. Usually the cortex is removed during the process of stone tool creation.
Ethnocentric: Applying negative judgments to other cultures based on comparison to one’s own.
Exogenous DNA: DNA that originates from sources outside of the specimen you are trying to sequence.
Flexed position: Fetal position, in which the legs are drawn up to the middle of the body and the arms are drawn toward the body centre. Intentional burials are often found in the flexed body position.
Foraminifera: Microscopic single-celled organisms with a shell that are common in all marine environments. The fossil record of foraminifera extends back well over 500 million years.
Glaciation: A glacial period, or time when a large portion of the world is covered by glaciers and ice sheets.
Globular: Round-shaped, like a globe.
Grave goods: Items included with a body at burial. Items may signify occupation or hobbies, social status, or level of importance in the community, or they may be items believed necessary for the afterlife.
Haft: A handle. Also used as a verb—to attach a handle to an item, such as a stone tool.
Infraorbital foramina: Small holes on the maxilla bone of the face that allows nerves and blood to reach the skin.
Insular dwarfing: A form of dwarfism that occurs when a limited geographic region, such as an island, causes a large-bodied animal to be selected for a smaller body size.
Interglacial: A warmer period between two glacial time periods.
Levallois technique: A distinctive technique of stone tool manufacturing used by Archaic Homo sapiens, including Neanderthals. The technique involves the preparation of a core and striking edges off in a regular fashion around the core. Then a series of similarly sized pieces can be removed, which can then be turned into different tools.
Midfacial prognathism: A forward projection of the nose or the middle facial region. Usually associated with Neanderthals.
Mousterian tools: The stone tool industry of Neanderthals and their contemporaries in Africa and Western Asia. Mousterian tools are known for a diverse set of flake tools, which is different from the large bifacial tools of the Acheulean industry.
Nasal aperture: The opening for the nose visible on a skull. Often pear- or heart-shaped.
Occipital bun: A prominent bulge or projection on the back of the skull, specifically the occipital bone. This is a feature present only on Neanderthal skulls.
Ochre: A natural clay pigment mixed with ferric oxide and clay and sand. Ranges in colour from brown to red to orange.
Retracted face: A face that is flatter.
Retromolar gap: A space behind the last molar and the end of the jaw. This is a feature present only on Neanderthals. It also occurs through cultural modification in modern humans who have had their third molars, or wisdom teeth, removed.
For Further Exploration
Anne and Bernard Spitzer Hall of Human Origins—American Museum of Natural History.
“Dawn of Humanity,” PBS documentary, 2015
“DNA Clues to Our Inner Neanderthal,” TED Talk by Svante Pääbo, 2011.
“The Dirt” Podcast, Episode 30, “The Human Family Tree (Shrub? Crabgrass? Tumbleweed?), Part 3: Very Humany Indeed”.
Frank, Rebecca. 2021. “The Genus Homo.” In Explorations: Lab and Activity Manual, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.
Hobbits on Flores, Indonesia - Smithsonian Human Origins.
Lumping or Splitting in the Fossil Record - UC Berkeley Understanding Evolution.
Neandertals and More - Max Planck Institute for Evolutionary Anthropology.
Neanderthals: Body of Evidence - SAPIENS.
Perash, Rose L., and Kristen A. Broehl. 2021. “Hominin Review: Evolutionary Trends.” In Explorations: Lab and Activity Manual, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.
Perkl, Bradley. “Brain, Language, Lithics.” In Explorations: Lab and Activity Manual, edited by. Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. CC BY-NC. Arlington, VA: American Anthropological Association.
Shanidar 3 - Neanderthal Skeleton - Smithsonian Human Origins.
Species - Smithsonian Human Origins.
Smithsonian Human Origins Program Facebook page (@smithsonian.humanorigins).
Paleoartist Brings Human Evolution to Life - Elisabeth Daynés.
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Acknowledgements
The authors would like to extend their thanks to Cassandra Gilmore and Anna Goldfield for thoughtful and insightful suggestions on the first edition of this chapter.
Katie Nelson, Ph.D., Inver Hills Community College
Lara Braff, Ph.D., Grossmont College
Beth Shook, Ph.D., California State University, Chico
Kelsie Aguilera, M.A., University of Hawai‘i: Leeward Community College
This chapter is a section of a revision from "Chapter 1: Introduction to Biological Anthropology” by Katie Nelson, Lara Braff, Beth Shook, and Kelsie Aguilera. In Explorations: An Open Invitation to Biological Anthropology, first edition, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff, which is licensed under CC BY-NC 4.0.
Learning Objectives
- Explain key components of the scientific method.
- Differentiate between hypotheses, theories, and laws.
- Differentiate science from other ways of knowing.
Anthropologists as Scientists
Biological anthropologists use the scientific method as a way of learning about the world around them. Many people think of science as taking place in a sterile laboratory, but in biological anthropology it is just as likely to occur somewhere else, such as at a research station in Ethiopia, a field site in Tanzania, or a town in El Salvador. To understand how information in this field is established, it is important to recognize what science is and is not, as well as to understand how the scientific method actually works.
Recognizing Science
Science combines our natural curiosity with our ability to experiment so we can understand the world around us and address needs in our communities. Thanks to science, meteorologists can predict the weather, it takes a relatively small number of farmers to grow enough food to feed our large population, our medicine continues to improve, and over half of the world’s population owns a cell phone.
Anyone can participate in science—not just academics. In fact, children are often some of the best scientists (Figure 1.15). An early, well-known psychologist, Jean Piaget (1896–1980), argued that a child is a “little scientist,” internally motivated to experiment and explore their world. This can be seen when an infant repeatedly drops a toy to see if the parent will pick it up, or when a four-year-old sincerely asks “why” again and again. Maria Montessori (1870–1952), an Italian doctor and educator, was interested in how children learn. Through her research, she also recognized that children have natural scientific tendencies. Children have a desire to explore their environment, ask questions, use their imaginations, and learn by doing. In 1907, Montessori opened a school to foster children’s natural desire to learn this way. This developed a child-centered teaching method that has spread around the world and is being used in over 22,000 schools today. In anthropology and other scientific fields, the process of learning is more formalized, but scientists still benefit from the curiosity that motivates children and still experience the thrill of discovery.

Science represents both a body of knowledge and the process for learning that knowledge (the scientific method). Scientific claims can, at times, be difficult to distinguish from other information. Science also incorporates a broad range of methods to collect data, adding to the difficulty of knowing what science really is. This section will address four key characteristics that help us define and recognize science: (1) science studies the physical and natural world and how it works, (2) scientific explanations must be testable and refutable, (3) science relies on empirical (observable) evidence, and (4) science involves the scientific community.
Science Studies the Physical and Natural World and How It Works
Our physical and natural universe ranges from very small (e.g., electrons) to very large (e.g., Earth itself and the galaxies beyond it). Scientists often design their research to address how and why natural forces influence our physical and natural world. In biological anthropology, we focus our questions on humans as well as other primate species, both living and extinct. We ask questions like: What influences a primate’s diet? Why do humans walk on two legs? And did Neanderthals and modern humans interbreed?
There are very few questions that are considered off-limits in science. That being said, the scope of scientific investigation is generally focused on natural phenomena and natural processes and excludes the supernatural. People often regard the supernatural, whether it be a ghost, luck, or god, as working outside the laws of the universe, which makes it difficult to study with a scientific approach. Science neither supports nor contradicts the existence of supernatural powers—it simply does not include the supernatural in its explanations.
Scientific Explanations Must Be Testable and Refutable
The goal of scientists is to identify a research question and then identify the best answer(s) to that question. For example, an excavation of a cemetery may reveal that many people buried there had unhealed fractures when they died, leading the anthropologist to ask: “Why did this population experience more broken bones than their neighbors?” There might be multiple explanations to address this question, such as a lack of calcium in their diets, participation in dangerous work, or violent conflict with neighbors; these explanations are considered hypotheses. In the past, you might have learned that a hypothesis is an “educated guess,” but in science, hypotheses are much more than that. A scientific hypothesis reflects a scientist’s knowledge-based experiences and background research. A hypothesis is better defined as an explanation of observed facts; hypotheses explain how and why observed phenomena are the way they are.
Scientific hypotheses should generate expectations that are testable. For example, if the best explanation regarding our cemetery population was that they were experiencing violent conflict with their neighbors, we should expect to find clues, like weapons or protective walls around their homes, in the anthropological record to support this. Alternatively, if this population did not experience violent conflict with their neighbors, we may eventually be able to gather enough evidence to rule out (refute) this explanation. An important part of science is rigorous testing. Science does not prove any hypothesis. However, a strong hypothesis is one that has strong supporting evidence and has not yet been disproven.
Science Relies on Empirical Evidence
The word empirical refers to experience that is verified by observation (rather than evidence that derives primarily from logic or theory). In anthropology, much evidence about our world is collected by observation through fieldwork or in a laboratory. The most reliable studies are based on accurately and precisely recorded observations. Scientists value studies that explain exactly what methods were used so that their data collection and analysis processes are reproducible. This allows for other scientists to expand the study or provide new insights into the observations.
Science Involves the Scientific Community
Contrary to many Hollywood science fiction films, good science is not carried out in isolation in a secret basement laboratory; rather, it is done as part of a community. Scientists pay attention to what others have done before them, present new ideas to each other, and publish in scientific journals. Most scientific research is collaborative, bringing together researchers with different types of specialized knowledge to work on a shared project. Today, thanks to technology, scientific projects can bring together researchers from different backgrounds, experiences, locations, and perspectives. Most big anthropological questions such as “Where did modern humans develop?,” “What genetic changes make us uniquely human?,” and “How did cooperative behavior evolve?” cannot be addressed with one simple study but are tested with different lines of evidence and by different scientists over time.
Working within a scientific community supports one of the most valuable aspects of science: that science is self-correcting. Science that is openly communicated with others allows for a system with checks and balances: competing explanations can be proposed and questionable studies can be reevaluated. Ultimately, the goal is that through science the best explanations will stand the test of time.
How Science Works: The Scientific Method
Most students have learned the scientific method as a simple linear, or perhaps circular, process (see, e.g., Figure 1.16). Typically, the process is said to begin with making observations about the natural world. This leads to the development of a scientific hypothesis. From the hypothesis a set of predictions can be made, which are then tested by experimentation or by making additional observations. Scientific predictions are often phrased as “if... then…” statements, such as “If hypothesis A is true, then this experiment will show outcome B.” The results of a scientific study should then either support or reject the hypothesis.

This simple version of the scientific method is valuable because it highlights the key aspects that should be present in any scientific research experiment or scientific paper. However, this simplistic view does not accurately represent the dynamic and creative side of science, nor does it identify the complex steps that are incorporated into a scientist’s routine.

Figure 1.17 provides an alternative representation of the scientific method that emphasizes the many paths to scientific discovery. While still incorporating the key components of making observations, testing ideas, and interpreting results, this chart shows that scientific ideas have many possible starting points and influences, and scientists often repeat steps and circle back around. Gathering evidence does not always rest on experiments in the laboratory. Evaluating data is not always clear-cut, and results are sometimes surprising or inconclusive. Many important discoveries were in fact made by mistake. For example, engineer Percy Spencer accidentally melted a chocolate bar in his pocket with a magnetron, which became the first microwave, and Spencer Silver invented the adhesive for 3M Post-it ® notes while trying to develop a strong glue. The real scientific process is more similar to the philosophy of the animated television character Ms. Frizzle from The Magic School Bus, “Take chances, make mistakes, get messy.”
Two key components lacking in the simple version of the scientific method are exploration and discovery. There are many reasons that a scientist might choose a particular research question: they may be motivated by personal experience, struck by something they read, or inspired by a student’s question in class. Often scientific research reveals more questions than answers, so experienced researchers rarely lack problems to solve. But identifying a research question is just part of the process; most scientists spend more time exploring the literature, sharing ideas, asking questions, and planning their research project than conducting the test itself.
Science itself is a social enterprise that is influenced by cultural issues and values, as well as funding priorities. For example, corporations are the biggest funders of scientific research, followed by government agencies such as the National Science Foundation (which also fund many research projects done at colleges and universities). Those organizations have great influence on what is considered valuable research at any given time. For example, the World Health Organization (WHO) has classified many diseases as “neglected tropical diseases,” including dengue, leprosy, rabies, and hookworm. Together these diseases affect an estimated one billion people, mostly in impoverished areas. While these debilitating tropical diseases can be as deadly as diseases that receive more attention, like AIDS and tuberculosis, they receive comparatively little funding due to political priorities (Farmer et al. 2013).
Also important to the scientific process are interactions within the scientific community. Scientific collaboration can take place through informal discussion over a cup of coffee as well as more formal interactions, such as presenting at conferences and engaging in scholarly peer review. Scholarly peer review describes the process whereby an author’s work must pass the scrutiny of other experts in the field before being accepted for publication in a journal or book. This helps keep scientists accountable for ethically responsible research projects and papers. Additionally, presenting data at conferences and in articles and books allows researchers to receive critical feedback from academic peers and others to test these ideas and further the field of science toward identifying the best explanations. It is important that the scientific field include researchers with diverse identities, backgrounds, and experiences so that researchers ask new questions, innovate, and problem solve more effectively.
Hypotheses, Theories, and Laws
Scientific investigation occurs at many levels, from investigating individual cases (e.g., “What is causing this child’s mysterious illness?”) to understanding processes that affect most of us (“What is the ideal amount of sleep for an adult?”). All of these questions are important and will generate different types of testable scientific explanations. So far, we have used the term hypothesis to describe these scientific ideas about why observed phenomena are the way they are. Hypotheses are typically explanations that address a narrow set of phenomena, such as (in anthropology) a particular human population or primate species.
In science, a theory is an explanation of observations that addresses a wide range of phenomena. Like hypotheses, theories also explain how or why something occurs, rely on empirical evidence, and are testable and able to be refuted. Because the term theory is often used casually outside of science, you may hear people try to dismiss a scientific claim as “just a theory.” In science there are often multiple competing theories, but over time some are eliminated, leaving standing the theory or theories that best explain the most evidence. Scientific theories that have stood the test of time are thus supported by many lines of evidence and are usually reliable. Some well-tested theories accepted by most scientists include the theory of general relativity, which explains the law of gravitation and its relation to other forces, and evolutionary theory, which describes how heritable traits can change in a population over time.
While scientific hypotheses and theories share many characteristics, laws are quite different. A law is a prediction about what will happen given certain conditions, not an explanation for how or why it happens. A law is not a “mature” version of a theory. For example, Newton’s universal law of gravity allows us to predict the gravitational force (F) between any two objects using the equation F=G(m1m2)/r2, but it does not explain why gravity works. Laws are often mathematical, and some well-known laws include Newton’s three laws of motion and Mendel’s laws of genetic inheritance. Laws are important, and their discovery often promotes the development of theories.

To demonstrate how important laws can be—and to show how unusual things can inspire scientific discoveries—we can use the story of the ancient Greek mathematician and inventor Archimedes (Figure 1.18). Archimedes’s buoyancy principle is a law that is useful for many things, including density calculations and designing ships. Purportedly, he made this discovery when he noticed the water level rise in the bathtub when he climbed in it. Realizing its importance, he is said to have shouted “Eureka” and proceeded to run naked through the city of Syracuse. While this fun story may or may not be true, it does remain that scientific laws, alongside scientific hypotheses and theories, have a very important role in the scientific process and in generating scientific explanations about our natural world.
Ways of Knowing: Science, Faith, and Anthropology
In anthropology, we recognize that there are many ways of knowing things. For instance, you might know that fingernails are softer than metal because as a child you accidentally stapled through your fingernail while doing an art project (a coauthor of this textbook once experienced this). This would be an example of knowledge you gained through experience. You might also know that inserting a knife into an electrical outlet is dangerous and could greatly harm you. Hopefully you learned this not from personal experience but through instruction from parents, teachers, and others in your social group. The degree to which humans rely on and benefit from the experiential knowledge of others is an important characteristic of what makes us human.
A unified way of knowing that is shared by a group of people and used to explain and predict phenomena is called a knowledge system. Human knowledge systems are diverse and reflect the wide range of cultures and societies throughout the world and through time.
Science and religion are both knowledge systems. Yet they differ in important ways. The type of knowledge gained from science is often called scientific understanding. As we have explored in the previous section, scientific understanding can change and relies on evidence and rigorous, repeated testing. Religious or spiritual ways of knowing are called belief, which is different from scientific understanding because they do not require repeated testing or validation (although they can rely on observations and experiences). Instead, belief relies on trust and faith.
Different individuals, cultures, and societies may place more value on one type of knowing than another, although most use a combination that includes science, empiricism, and religion. In fact, Bronisław Malinowski (1884–1942), an important anthropologist of the early twentieth century, concluded that all societies use both religion and science in some way or another, because they are both common ways that humans experience the world.
In contemporary societies such as the United States, science and (some) religions conflict on the topic of human origins. Nearly every culture and society has a unique origin story that explains where they came from and how they came to be who they are today. These stories are often integrated into the culture’s religious belief system. Many anthropologists are interested in faith-based origin stories and other beliefs because they show us how a particular group of people explain the world and their place in it. Anthropologists also value scientific understanding as the basis for how humans vary biologically and change over time. In other words, anthropologists value the multiple knowledge systems of different groups and use them to understand the human condition in a broad and inclusive way.
It is also important to note that scientists often depend on the local knowledge of the people with whom they work to understand elements of the natural or physical world that science has not yet investigated. Many groups, including Indigenous peoples, know about the world through prolonged relationships with the environment. Indigenous knowledge systems—specific to an Indigenous community or group—are informed by their own empirical observation of a specific environment and passed down over generations.
While religion and Indigenous knowledge systems may play a complementary role in helping anthropologists understand the human condition, they are distinct from science. The anthropological subdiscipline of biological anthropology is based on scientific ways of knowing about humans and human origins. In this volume, we will exclusively explore what science tells us about how humans came to be and why we are the way we are today. Therefore, you do not need to believe in evolution to master this material, because belief is not a scientific way of knowing. For this textbook, you only need to understand the scientific perspectives of evolution.
Throughout our lives, each of us work to reconcile our worldview with the different ways we have of knowing things. This is part of our lifelong intellectual journey. It is also, in our opinion, one of the most exciting parts of learning. We are pleased you have joined us on this journey of knowledge about humanity and yourself!
Summary
Biological anthropology relies on the scientific method to investigate humans and other primates across time and place. While research in this field begins with careful observation, hypotheses are more closely examined with empirical evidence from excavation, fieldwork, or laboratory study. Focusing on this scientific foundation, the discipline allows anthropologists to move beyond belief and speculation to produce knowledge that is testable and continually open to challenge.
Findings are continually debated, reviewed, and revised, ensuring that explanations improve and remain grounded in current and relevant evidence. If you have not caught on already, this textbook follows the same process, reflecting the evolving body of knowledge that is biological anthropology.
Review Questions
- What is science?
- What is the scientific method?
- How does science compare to other ways of knowing?
Key Terms
Empirical: Evidence that is verifiable by observation or experience instead of relying primarily on logic or theory.
Faith: Complete trust or confidence in the doctrines of a religion, typically based on spiritual apprehension rather than empirical proof.
Knowledge system: A unified way of knowing that is shared by a group of people and used to explain and predict phenomena.
Participant observation: A research method common in cultural anthropology that involves living with, observing, and participating in the same activities as the people one studies.
Scholarly peer review: The process whereby an author’s work must pass the scrutiny of other experts in the field before being published in a journal or book.
Scientific understanding: Knowledge accumulated by systematic scientific study, supported by rigorous testing and organized by general principles.
Theory: An explanation of observations that typically addresses a wide range of phenomena.
For Further Exploration
Understanding Science website (a project of the University of California Museum of Paleontology.
Anticole, Matt. n.d. “What’s the Difference between a Scientific Law and Theory?" TedEd Lesson. Accessed January 28, 2023. 28, 2023.
Chan, Keith. 2021. “Icebreaker Science.” In Explorations: Lab and Activities Manual, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.
Chizmeshya, Sydney Quinn, and Katherine E. Brent. 2021. “Knowing and Believing.” In Explorations: Lab and Activities Manual, edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.
Pfister, Anne E. 2021. “Science and Belief: Just Because We Can, Doesn’t Always Mean We Should.” In Explorations: Lab and Activities Manual edited by Beth Shook, Katie Nelson, Kelsie Aguilera, and Lara Braff. Arlington, VA: American Anthropological Association.
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